A review of the use of pteridophytes for treating human ailments

A review of the use of pteridophytes for treating human ailments 蕨类植物治疗人类疾病的应用概况 1 2 first-author 3 2 2 1 corresp ‡Corresponding Author ‡Corresponding Author †E-mail:lsslwb@mail.sysu.edu.cn †E-mail:lsslwb@mail.sysu.edu.cn lsslwb@mail.sysu.edu.cn State Key Laboratory of Biocontrol and Guangdong Key Laboratory of Plant Resources, School of Life Sciences, Sun Yat-sen University, Guangzhou 510275, China State Key Laboratory of Biocontrol and Guangdong Key Laboratory of Plant Resources, School of Life Sciences, Sun Yat-sen University, Guangzhou 510275, China Shenzhen Key Laboratory of Southern Subtropical Plant Diversity, Fairy Lake Botanical Garden /Chinese Academy of Sciences, Shenzhen 518004, China Shenzhen Key Laboratory of Southern Subtropical Plant Diversity, Fairy Lake Botanical Garden /Chinese Academy of Sciences, Shenzhen 518004, China Department of Zoology, St. Xavier’s College, Palayamkottai, Tamil Nadu 627002, India Department of Zoology, St. Xavier’s College, Palayamkottai, Tamil Nadu 627002, India The aim of this review was to explore the pharmacological activity of early tracheophytes (pteridophytes) as an alternative medicine for treating human ailments. As the first vascular plants, pteridophytes (aka, ferns and fern allies) are an ancient lineage, and human beings have been exploring and using taxa from this lineage for over 2000 years because of their beneficial properties. We have documented the medicinal uses of pteridophytes belonging to thirty different families. The lycophyte Selaginella sp. was shown in earlier studies to have multiple pharmacological activity, such as antioxidant, anti-inflammatory, anti-cancer, antidiabetic, antiviral, antimicrobial, and anti-Alzheimer properties. Among all the pteridophytes examined, taxa from the Pteridaceae, Polypodiaceae, and Adiantaceae exhibited significant medicinal activity. Based on our review, many pteridophytes have properties that could be used in alternative medicine for treatment of various human illnesses. Biotechnological tools can be used to preserve and even improve their bioactive molecules for the preparation of medicines against illness. Even though several studies have reported medicinal uses of ferns, the possible bioactive compounds of several pteridophytes have not been identified. Furthermore, their optimal dosage level and treatment strategies still need to be determined. Finally, the future direction of pteridophyte research is discussed. 蕨类植物全世界约48科250属12 000种,目前只有少数蕨类植物的生物学活性得到了研究,这些研究已经证明蕨类植物的药用价值及其在植物科学领域的重要性.本文旨在总结关于蕨类植物的主要化学成分及药理活性,特别是其抗氧化活性、抗肿瘤活性、抗糖尿病活性、抗病毒活性、抗炎活性、抗菌活性、抗老年痴呆症和脑疾病活性等,可用于替代医学的各种人类疾病治疗.然而,它们的最佳剂量和治疗策略仍需要进一步研究. Pteridophyte Fern Alternative medicine Selaginella Human illness Bioactive molecule 蕨类植物 替代医学 卷柏属 人类疾病 生物活性分子 Introduction 1 Medicinal plants have been the source of remedies for the treatment of several illnesses since earliest times, and people from all continents have practiced the use of botanicals for millenia. Despite the significant growth in synthetic organic chemistry in the 20th century, over 25% of approved drugs in industrialized countries are obtained directly or indirectly from plants (Newman et al., 2000 2000 ). In recent years, the search for phytochemicals with antioxidant, antimicrobial, or anti-inflammatory properties has been on the rise due to their potential use for the treatment of various chronic and infectious diseases (Halliwell, 1996 1996 ). Therapeutic plants may be an alternative source of antimicrobial agents with important bioactivity against pathogenic and transmissible microorganisms, with fewer side effects than synthetic antibiotics (Berahou et al., 2007 2007 ). Plant extracts have numerous secondary metabolites, such as phenolic components, that enhance biological activity. Plant secondary metabolites are known to possess antimicrobial and antioxidant properties, and some of them are categorized as generally recognized safe substances (Proestos et al., 2005 2005 ). With the growing interest in these compounds, phytocompounds from therapeutic plants with some biomedicinal activity are in great demand (Chen et al., 2008 2008 ; Pesewu et al., 2008 2008 ; Prachayasittikul et al., 2008 2008 ). Ferns and their allies are one of the oldest major divisions of the Pteridophyta, and comprise over 12 000 species spread among 250 different genera (Chang et al., 2011 2011 ). Traditionally, the biomedical system and Ayurvedic systems of medicine, named Sushruta (ca. 100 AD) and Charka (ca. 100 AD), suggested the use of some ferns in the Samhita texts. Pteridophytes are also used by physicians in the Unani system of medicine (Uddin et al., 1998 1998 ). In the traditional Chinese system of medicine, several ferns are recommended by native doctors (Kimura and Noro, 1965 1965 ). In more recent times, ethnobotanical and advanced pharmacological studies have been carried out on ferns and their allies by several investigators (Dhiman, 1998 1998 ; Vasudeva, 1999 1999 ; Reddy et al., 2001 2001 ; Singh L et al., 2001 2001 ; Gogoi, 2002 2002 ; Singh M et al., 2008a 2008a , 2008b 2008b ; Chen JJ et al., 2005 2005 ; Chen K et al., 2005 2005 ). Phytochemicals of ferns and their distribution in other plant groups 1 The secondary metabolites, flavonoids, have anti-inflammatory properties through the inhibition of the cyclo oxygenase (COX) pathway (Liang et al., 1999 1999 ). They were also found to have antioxidant, anti-cancer, and antimicrobial properties (Lin et al., 2008 2008 ; Yoshida et al., 2008 2008 ; Amaral et al., 2009 2009 ; Lopez-Lazaro, 2009 2009 ; Govindappa et al., 2011 2011 ). Spider brake fern ( Pteris multifida ) has various bioactive flavonoids with heat-clearing, antipyretic, detoxification, antibiotic, anti-inflammatory, and antimutagenic activity (Lee and Lin, 1988 1988 ). The entire plant of P. multifida contains various bioactive compounds including: luteolin-7- O -glucoside (Murakami and Machashi, 1985 1985 ; Lu et al., 1999 1999 ), 16-hydroxy-kaurane-2-β- D -glucoside (Liu and Qin, 2002 2002 ), luteolin, palmitic acid, apigenin 4- O -α- L -rhamnoside (Lu et al., 1999 1999 ; Qin et al., 2006 2006 ), quercetin, hyperin, isoquercitrin, kaempferol, rutin (Lu et al., 1999 1999 ; Wang et al., 2010 2010 ; Hoang and Tran, 2014 2014 ), and apigenin-7- O -β- D -glucoside (Lu et al., 1999 1999 ; Hoang and Tran, 2014 2014 ). Also, the cytotoxic bioactive compounds, pterosin sesquiterpenes, have been isolated from P. multifida (Shu et al., 2012 2012 ). Bioactive terpenoids, especially the monocyclic sesquiterpene α-caryophyllene, were found in an ethanolic extract of Pteris tripartita (Baskaran and Jeyachandran, 2010 2010 ). Sesquiterpenes form a significant group of secondary metabolites and have a varied range of medicinal activity, including anti-cancer, anti-inflammatory, cytotoxic, plant growth-regulatory, and antimicrobial properties (Baruah et al., 1994 1994 ). The anti-inflammatory activity of fatty acids such as myristic, palmitic, stearic, oleic-linoleic, and eicosatrienoic acids are widely known (Bremner and Heinrich, 2002 2002 ). Stearic acid has been shown to have both anti-inflammatory and hepatoprotective effects (Pan et al., 2010 2010 ). Baskaran and Jeyachandran ( 2010 2010 ) found octadecanoic acid (a stearic acid) in an ethanol extract of P. tripartita . Pteridium aquilinum , thought to be a fern with potent anti-cancer properties, contains p -coumaric, p -hydroxybenzoic, caffeic, ferulic, vanillic, and protocatechuic acids (Michael and Gillian, 1984 1984 ). Hydroxycinnamic acid, anthocyanins, total polyphenols, and flavonoids have been quantified in Stenochlaena palustris (Chai et al., 2012 2012 ). Some bioactive flavones such as amentoflavone, robustaflavone, biapigenin, hinokiflavone, podocarpusflavone A, and ginkgetin, in Selaginella sp. were reported to have antioxidant, antivirus, and anti-cancer activity (Ma et al., 2001 2001 ; Shi et al., 2008 2008 ; Liu et al., 2011 2011 ; Li S et al., 2014 2014 ). Both the whole plant and mature spores of Lygodium japonicum are used for medical purposes in traditional Chinese medicine. For over 2000 years, L. japonicum has been used in China for the treatment of gorge gall, skin eczema, enteritis diarrhea, nephritis dropsy, urine calculus, and infection (Wynne et al., 1998 1998 ). The foremost phenolic components, namely 4-hydroxycinnamic acid ( p- coumaric), 3-methoxy-4-hydroxycinnamic acid (ferulic), 3,4-dihydroxycinnamic acid (caffeic), 3-methoxy-4-hydroxybenzoic, acid and 3-caffeoilquinic acid (chlorogenic), have been found in Polypodium leucotomos extracts (Gombau et al., 2006 2006 ). The free radical scavenging bioactive principles viz. flavan-4-ol glycosides, abacopterins, huperzine A, isoquercetin, di- E -caffeoyl- meso -tartaric acid, flavaspidic acid PB, flavaspidic acid AB, flavan-3-ol, kaempferol, A-type proanthocyanidins, and afzelechin, have been derived from a few ferns including Abacopteris penangiana , Huperzia selago , Equisetum arvense , and Dryopteris crassirhizoma (Lee et al., 2003 2003 ; Staerk et al., 2004 2004 ; Zhao et al., 2007 2007 ; Mimica et al., 2008 2008 ). Fig. 1 1 and Table 1 1 list the distribution of numerous bioactive compounds present in pteridophytes. No. Pteridophyte Bioactive compounds Source(s) 1   Pteris multifida  Luteolin-7- O -glucoside, 16-hydroxy-kaurane-2-β- D -glucoside, luteolin, palmitic acid, apigenin4- O -α- L -rhamnoside, quercetin, hyperin, isoquercitrin, kaempferol, rutin, apigenin-7- O -β- D -glucoside, and pterosin  Murakami and Machashi, 1985 1985 ; Lu et al., 1999 1999 ; Liu and Qin, 2002 2002 ; Qin et al., 2006 2006 ; Wang et al., 2010 2010 ; Shu et al., 2012 2012 ; Hoang and Tran, 2014 2014 2 Pteris tripartita  α-Caryophyllene and octadecanoic acid  Baskaran and Jeyachandran, 2010 2010 3 Abacopteris penangiana  Flavan-4-ol, glycosides, and abacopterins  Zhao et al., 2007 2007 4 Pteridium aquilinum p -Coumaric acid, p -hydroxybenzoic acid, caffeic acid, ferulic acid, vanillic acid, protocatechuic acid, kaempferol, quercetin, and apigenin  Michael and Gillian, 1984 1984 5 Selaginella sp.  Amentoflavone, robustaflavone, biapigenin, hinokiflavone, podocarpusflavone A, and ginkgetin  Ma et al., 2001 2001 ; Shi et al., 2008 2008 ; Liu et al., 2011 2011 ; Li S et al., 2014 2014 6 Equisetum arvense  Isoquercetin, quercetin 3- O -glucoside, quercetin 3- O -(6''- O -malonylglucoside), 5- O -caffeoyl meso -tartaric acid, monocaffeoyl meso -tartaris acid, monocaffeoyl meso -tartaris acid, di- E -caffeoyl- meso -tartaric acid, hexahydrofarnesyl acetone, cis -geranyl scetone, thymol, and trans -phytol  Radulovic et al., 2006 2006 ; Milovanovic et al., 2007 2007 ; Mimica et al., 2008 2008 7 Polypodium leucotomos  4-Hydroxycinnamic acid ( p -coumaric), 3-methoxy-4-hydroxycinnamic acid (ferulic), 3,4-dihydroxycinnamic acid (caffeic), 3-methoxy-4-hydroxybenzoic acid, and 3-caffeoilquinic acid (chlorogenic)  Gombau et al., 2006 2006 8 Adiantum capillus-veneris  Adiantone, adiantoxide, astragalin, β-sitosterol, caffeic acids, caffeylgalactose, caffeylglucose, campesterol, carotenes, coumaric acids, coumarylglucoses, diplopterol, epoxyfilicane, fernadiene, fernene, filicanes, hopanone, hydroxy-adiantone, hydroxy-cinnamic acid, isoadiantone, isoquercetin, kaempferols, lutein, mutatoxanthin, naringin, neoxanthin, nicotiflorin, oleananes, populnin, procyanidin, prodelphinidin, quercetins, querciturone, quinic acid, rhodoxanthin, rutin, shikimic acid, violaxanthin, and zeaxanthin  Taylor, 2003 2003 9 Dryopteris crassirhizoma  Flaspidic acid PB and flavaspidic acid AB  Lee et al., 2003 2003 10 Cyathea gigantea  β-Sitosterol  Woyengo, 2009 11 Cibotium barometz  Pterosins, terpenoids, sterides, flavones, glucosides, aromatic, and pyrone compounds  Wu et al., 2007 2007 ; Xu et al., 2012 2012 12 Cyathea phalerata  Kaempferol-3-neohesperidoside, 4- O -β- D -glucopyranosyl caffeic acid, 4- O -β- D -glucopyranosyl p -coumaric acid, 3,4-spyroglucopyranosyl protocatechuic acid, sitosterol β- D -glucoside, β-sitosterol, kaempferol, and vitexin  Hort et al., 2008 2008 13 Pteris vittata  Rutin, kaempferol monoglycoside, kaempferol diglycoside, quercetin monoglycoside, and quercetin diglycoside  Salatino and Prado, 1998 1998 ; Singh et al., 2008b 2008b 14 Polypodium leucotomos  3,4-Dihydroxybenzoic acid, chlorogenic acid, 4-hydroxybenzoic acid, chlorogenic acid, vanillic acid, caffeic acid, 4-hydroxycinnamic acid, 4-hydroxycinnamoyl-quinic acid, and ferulic acid  Garcia et al., 2006 2006 15 Angiopteris evecta  Angiopterioside  Twentyman et al., 1989 1989 16 Onoclea sensibilis  Kaempferol and quercetin  Harada and Saiki, 1955 1955 17 Matteuccia struthiopteris  Kaempferol and quercetin 18 Equisetum ramoissimum  Apigenin and luteolin 19   Lycopodium japonicum  Lycopodine-type alkaloids  He et al., 2014 20   Lycopodium sieboldii  Sieboldine A  Hirasawa et al., 2003 21   Lycopodium casuarinoides  Huperzine B and N -demethylhuperzinine  Shen and Chen, 1994 22   Selaginella tamariscina  Caffeic acid, ferulic acid, vanillic acid, syringic acid, adenosine, umbelliferone, and amentoflavone  Bi et al., 2004; Zheng et al., 2004; Woo et al., 2005 23   Selaginella moellendorfii  Moellenoside B and selaginellic acid  Wang et al., 2009b; Feng et al., 2011 24   Selaginella doederleinii  Apigenin and hordenine  Chao et al., 1987; Chen et al., 1995 25   Dicranopteris linearis  Afzelin, quercitrin, isoquercitrin, astragarin, rutin, kaempferol 3- O -(4- O - p -coumaroyl-3- O -α-lrhamnopyranosyl)-α- L -rhamnopyranosyl-(1→6)-β- D -glucopyranoside, and (6S,13S)-6-[6- O -acetyl-β- D -glucopyranosyl-(1→4)-α- L -rhamnopyranosyloxy]-13-[α-lrhamnopyranosyl-(1→4)-β- D -fucopyranosyloxy]cleroda-3,14-diene  Raja et al., 1995; Zakaria et al., 2008 26   Psilotum nudum  Quercetin, kaempferol, amentoflavone, hinokiflavone, vicenin-2 psilotin, and 3'-hydroxypsilotin  Cambie and Ash, 1994 27   Huperzia selago  Huperzine A  Staerk et al., 2004 28   Huperzia serrata  Alkaloids: huperzine A and B  Liu et al., 1986 29   Pityrogramma tartarea  2'6'-dihydroxy-4,4'-dimethoxydihydrochalcone, kaempferol 7-methyl ether, and apigenin 7-methyl ether  Star and Mabry, 1971 30   Pityrogramma calomelanos 31   Cheilanthes concolor  7- O -Glycosides of apigenin, aglycone, 7- O -glycosides of luteolin, 7- O -glycosides of chrysoeriol, 3- O -glycosides of kaempferol, 3- O -glycosides of kaempferol, and 3- O -glycosides of quercetin  Salatino and Prado, 1998 32   Cheilanthes flexuosa 33   Cheilanthes goyazensis 34   Doryopteris ornithopus 35   Pellaea cymbiformis 36   Pellaea gleichenioides 37   Pellaea pinnata 38   Pellaea riedelii 39   Pteris altissima 40   Pteris angustata 41   Pteris decurrens 42   Pteris deflexa 43   Pteris denticulata 44   Pteris plumula 45   Pteris podophylla 46   Pteris splendens 47   Pteris propinqua 48   Cyrtomium fortunei  Protocate chaldehyde, woodwardinsaure methylester, physcion, pimpinellin, ursolic acid, sitost-4-en-3-one, betulin, 3',4',5-trihydroxy-3,7-dimethoxyflavone, woodwardinic acid, sutchuenoside A, kaempferol-3,7- O -α- L -dirhamnoside, (−)-epicatechin, (+)-catechin hydrate kaempferol, crassirhizomoside A, and kaempferol-3- O -(3- O -acetyl-α- L -rhamnopyranoside  Yang et al., 2013 Marchantiophyta have a cellular oil body which produces numerous mono-, sesqui-, and di-terpenoids, aromatic compounds like bibenzyl, bis-bibenzyls, and acetogenins. Most sesqui-and di-terpenoids isolated from non-vascular plants (liverworts) are enantiomers of those present in vascular plants (Asakawa et al., 2013 2013 ). Since 1960, studies have begun to identify triterpenoids in ferns too. Moreover, flavonoids are present in more than one genus of ferns and also occur in fern allies (Berti and Bottari, 1968 1968 ). Pteridophytes are known to produce proanthocyanidins which are present in a polymerized molecular form, as tannins. Proanthocyanidins in higher groups of plants are formed by dihydroflavonol-4-reductase (DFR) using dihydroflavonols as substrate. The DFR enzyme has not been described in pteridophytes, but the genomic characterization of ferns has begun. DFR has been obtained from duplicate genes linked with primary metabolism; in this instance, nicotinamide adenine dinucleotide phosphate (NADPH)-dependent reductases are related to steroid metabolism (Baker and Blasco, 1992 1992 ). Moreover, three divisions of the flavonoid pathway well-known in the bryophytes are also found in pteridophytes including procyanidins, prodelphinidins, and flavonols like kaempferol, quercetin, and myricetin (Markham, 1988 1988 ). Acylphloroglucinol compounds are found only in the genus Dryopteris , which has been used in chemosystematic and phylogenetic studies (Widen, 1971 1971 ; Widen and Britton, 1971a 1971a , 1971b 1971b ). Flavonoids and phenolic compounds are the most general group of bioactive principles in plants, occurring in almost all plant parts, but mainly in photosynthesizing cells. These compounds are the main producers of the colors of blooming plants (Koes et al., 2005 2005 ). In early studies, myricetin, quercetin, kaempferol, apigenin, luteolin, and gallic acid were reported in angiosperms including the Hamamelidae, Rutaceae, Eriocaulaceae, Lythraceae, and Velloziaceae (Grieve and Scora, 1980 1980 ; Giannasi, 1986 1986 ; Salatino et al., 2000 2000 ). Several phenolic compounds, such as p -coumaric acid, caffeic acid, ferulic acid, protocatechuic acid, gallic acid, vanillic acid, and flavonoids like luteolin, apigenin, have also been identified in seagrasses (Subhashini et al., 2013 2013 ). Apigenin, kaempferol, luteolin, myricetin, quercetin, and rutin were identified in some Iranian flowering plants from different families including the Leguminosae, Polygonaceae, Euphorbiaceae, Resedaceae, and Cyperaceae (Mitra, 2012 2012 ). The bioactive flavonoids, kaempferol, luteolin, quercetin, quercitrin, isoquercitrin, hyperoside, and guaijaverin, have been derived from methanol extracts of Hypericum brasiliense (Rocha et al., 1995 1995 ). In addition, kaempferol, myricetin, catechin, vitexin, orientin, isoquercitin, hyperside, isovitexin, luteolin-7-glucoside, rutin, kampferol-7-neohesp-eridoside, quercitin, luteolin, and apigenin have been identified in the angiosperm families Lytheraceae, Acanthaceae, Chenopodiaceae, Caprifoliaceae, Euphorbiaceae, Verbenaceae, Zygophyllaceae, Moraceae, Asteraceae, Urticaceae, Scrophulariaceae, Verbenaceae, and Mimosaceae (Khan et al., 2012 2012 ). The evolution of flavonoids in land plants has been discussed from an enzymatic point of view. In ferns and their allies, proanthocyanidin and NADPH reductase are present for flavonoid synthesis while anthocyanidin and pterocarpan synthase have been reported in gymnosperms and angiosperms. Moreover, at two different levels, chalcone synthase, chalcone isomerase, anthocyanidin synthase, flavonol synthase, isoflavone synthase, and NADPH reductase are also involved in flavonoid synthesis (Stafford, 1991 1991 ). These studies show that all land plants are related based on the evidence of shared secondary metabolites. The genetic relationships among genes in response to phytochemical synthesis in vascular and non-vascular plant groups should be studied further to understand their phytochemical evolution. Activity of ferns 1 Antioxidant activity 2 Most of the pteridophytes are of great economic value, especially in medical, aesthetic, and food applications (Vasudeva, 1999 1999 ). Table 2 2 summarizes the antioxidant activity of pteridophytes. The antioxidant activity has been reported for several fern species, namely Polystichum semifertile , Nothoperanema hendersonii , Braomea insignis (Chen K et al., 2005 2005 ), some Chinese ferns (31 species) (Ding et al., 2008 2008 ), Selaginella sp. (Gayathri et al., 2005 2005 ), Polyopodium leucotomus (Garcia et al., 2006 2006 ; Gombau et al., 2006 2006 ), E . arvense , Equesitum sylvaticum (Milovanovic et al., 2007 2007 ), A . penangiana (Zhao et al., 2007 2007 ), Drynaria fortunei , Pseudodrynaria coronans , Davallia divaricata , Davallia mariesii , Davallia solida , Humata griffithiana (Chang et al., 2007b 2007b ), Marsilea quadrifolia (Fig. 2c 2c ) (Ripa et al., 2009 2009 ), Cheilanthes anceps Swartz (Chowdhary et al., 2010 2010 ), Pteris ensiformis Burm. (Chen et al., 2007 2007 ; Wei et al., 2007 2007 ), P . multifida (Shyur et al., 2005 2005 ; Wang et al., 2009a 2009a ), P. tripartita (Baskaran and Jeyachandran, 2010 2010 ), Dicranopteris linearis (Fig. 2d 2d ) (Zakaria et al., 2011 2011 ), and Drynaria quercifolia (Fig. 2e 2e ) (Milan et al., 2013 2013 ). Efficient antioxidant properties (0.02–0.12 mg/ml in 2,2-diphenyl-1-picrylhydrazyl (DPPH) activity and 0.03–0.22 mg/ml in activity of 2,2'-azino-bis-(3-ethyl benzthiazoline-6-sulphonie acid diammonium salt (ABTS) radical scavenging) of several fern genera like Davallia , Hypolepis , Pteridium , Cyrtominum , Dryopteris , Polystichum , Dicranopteris , Lycopodium , Osmunda , Adiantum , Coniogramme , Polypodium , Pyrrosia , Pteris , Lygodium , Selaginella , Thelypteris , Athyrium , Matteuccia , Onoclea , and Woodsia have also been reported (Shin, 2010 2010 ). According to Shin and Lee ( 2010 2010 ), scavenging effects on DPPH and ABTS radicals were higher in methanol extracts from rhizomes than in those from aerial parts of Adiantum pedatum , Coniogramme japonica , D . mariesii , Cyrtomium fortunei , D . crassirhizoma , Polystichum polyblepharum , Polystichum lepidocaulon , Dryopteris nipponensis , Pteris cretica , P. multifida , and Polypodium nipponica . Note: for interpretation of the references to color in this figure legend, the reader is referred to the web version of this article No. Plant name Family name Methods used Source(s) 1 Polystichum semifertile Dryopteridaceae 3 Xanthine oxidase, lipoxygenase activity, and ABTS •+ assay 3 Chen et al., 2005a 2 Nothoperanema hendersonii Dryopteridaceae 3 Braomea insignis Blechnaceae 4 Blechnum orientale Blechnaceae DPPH assay Lai et al., 2010 2010 5 Selaginella sp. ( n =3) Selaginellaceae In vitro lipid peroxidation, hydroxyl radical scavenging activity Gayathri et al., 2005 2005 6 Polypodium leucotomus Polypodiaceae H 2 O 2 and FRAP assays Garcia et al., 2006 2006 ; Gombau et al., 2006 2006 7 Equesitum sp. ( n =5) Equisetaceae Total antioxidant capacity Milovanovic et al., 2007 2007 8 Abacopteris penangiana Thelypteridaceae Trolox equivalent antioxidant capacity (TEAC) assay Zhao et al., 2007 2007 9 Marsilea quadrifolia Marsileaceae DPPH assay Ripa et al., 2009 2009 10 Cheilanthes anceps Sinopteridaceae DPPH assay Chowdhary et al., 2010 2010 11 Pteris ensiformis Pteridaceae DPPH and TEAC assays Chen et al., 2007 2007 ; Wei et al., 2007 2007 12 Pteris multifida Pteridaceae DPPH, superoxide scavenging activity Shyur et al., 2005 2005 ; Wang et al., 2009a 2009a 13 Pteris tripartita Pteridaceae DPPH, metal chelating activity, phosphomolydenum assay Baskaran and Jeyachandran, 2010 2010 14 Dicranopteris linearis Gleicheniaceae DPPH assay Milan et al., 2013 2013 15 Selaginella labordei Selaginellaceae In vitro xanthine oxidase inhibition Tan et al., 2009 2009 16 Diplazium polypodioides Athyriaceae DPPH assay Kshirsagar and Upadhyay, 2009 2009 17 Pteris vittata Pteridaceae 10 DPPH and ferric reducing power assays 10 Lai and Lim, 2011 2011 18 Adiantum radianum Pteridaceae 19 Acrostichum aureum Pteridaceae 20 Cyathea latebrosa Cyatheaceae 21 Cibotium barometz Dicksoniaceae 22 Drynaria quercifolia Polypodiaceae 23 Blechnum orientale Blechnaceae 24 Diplazium esculentum Athyriaceae 25 Nephrolepis biserrata Nephrolepidaceae 26 Dicranopteris linearis Gleicheniaceae 27 Dicksonia sellowiana Dicksoniaceae In vitro and in vivo antioxidant assays Bora et al., 2005 2005 ; Rattmann et al., 2011 2011 ; Oliveira et al., 2016 2016 28 Microgramma vacciniifolia Polypodiaceae In vitro DPPH assay Peres et al., 2009 2009 29 Polypodium leucotomos Polypodiaceae In vitro H 2 O 2 assay Gombau et al., 2006 2006 30 Davallia mariesii Davalliaceae 9 In vitro DPPH, ABTS •+ , and total antioxidant assays 9 Shin, 2010 2010 ; Delong et al., 2011 2011 31 Cyrtomium fortunei Dryopteridaceae 32 Dryopteris crassirhizoma Dryopteridaceae 33 Dryopteris nipponensis Dryopteridaceae 34 Polystichum lepidocaulon Dryopteridaceae 35 Polystichum polyblepharum Dryopteridaceae 36 Dicranopteris pedata Gleicheniaceae 37 Osmunda cinnamomea var. fokiensis Osmundaceae 38 Osmunda japonica Osmundaceae 39 Adiantum pedatum Pteridaceae 7 In vitro DPPH, ABTS •+ , and total antioxidant assays 7 Shin, 2010; Delong et al., 2011 40 Pyrrosia lingua Polypodiaceae 41 Lygodium japonicum Lygodiaceae 42 Thelypteris acuminate Thelypteridaceae 43 Athrium niponicum Athyriaceae 44 Onoclea sensibilis var. interrupta Onocleaceae 45 Matteuccia struthiopteris Onocleaceae 46 Aleuritopteris flava Pteridaceae 8 DPPH, ABTS •+ , ferric reducing power, and metal chelating assays 8 Gupta et al., 2014 47 Pteris scabristipes Pteridaceae 48 Adiantum edgeworthii Pteridaceae 49 Lindsaea odorata Dennstaedtiaceae 50 Microlepia rhomboidea Dennstaedtiaceae 51 Microlepia hallbergii Dennstaedtiaceae 52 Asplenium khasianum Aspleniaceae 53 Diplazium esculentum Athyriaceae 54 Athyrium filix-femina Athyriaceae 3 Oxygen radical absorbance capacity 3 Soare et al., 2012a 55 Dryopteris affinis Dryopteridaceae 56 Dryopteris filix-mas Dryopteridaceae 57 Dryopteris cochleata Dryopteridaceae DPPH, ABTS •+ , superoxide, nitric oxide, hydroxide, reducing power, ferrous chelating, and lipid peroxidation assays Kathirvel et al., 2014 According to Kshirsagar and Upadhyay ( 2009 2009 ), methanol extracts of Diplazium polypodioides and ethyl acetate or water extracts of Blechnum orientale show strong antioxidant capacity (Lai et al., 2010 2010 ). Lai and Lim ( 2011 2011 ) reported that Pteris vittata , Cyathea latebrosa , Cibotium barometz (Fig. 2f 2f ), D . quercifolia , B . orientale (Fig. 2g 2g ), Adiantum radianum , Diplazium esculentum , Acrostichum aureum , Nephrolepis biserrata , and D . linearis exhibited strong antioxidant activity ranging from 0.12 to 0.57 mg/ml. The presence of phenolics in extracts of Dicksonia sellowiana may contribute directly to its antioxidant properties (Duh et al., 1999 1999 ; Bora et al., 2005 2005 ). Ethanolic and ethyl acetate extracts showed good antioxidant action based on the DPPH radical scavenging assay in Microgramma vacciniifolia (Peres et al., 2009 2009 ). According to Hort et al. ( 2008 2008 ), the antioxidant activity and hepatoprotective potential of crude extracts are based on organic solvents of increasing polarity. Gombau et al. ( 2006 2006 ) demonstrated that hexane and chloroform extracts of P . leucotomos had significant inhibitory activity/strong antioxidative properties, based on an H 2 O 2 (hydrogen peroxide) assay. The antioxidant properties of methanol extracts of rhizomes and fronds of D . mariesii , C . fortunei , D . crassirhizoma , D. nipponensis , P . lepidocaulon , P. polyblepharum , Dicranopteris pedata , Osmunda cinnamomea var. fokiensis (Fig. 3f 3f ), Osmunda japonica , A . pedatum , Pyrrosia lingua , L . japonicum (Fig. 2b 2b ), Thelypteris acuminata , Athyrium niponicum , Matteuccia struthiopteris (Fig. 3h 3h ), and Onoclea sensibilis var. interrupta were reported by Shin ( 2010 2010 ) and Delong et al. ( 2011 2011 ). Note: for interpretation of the references to color in this figure legend, the reader is referred to the web version of this article Methanol leaf extracts of P . multifida showed stronger antioxidant activity than ethanol extracts (Hoang and Tran, 2014 2014 ). In a DPPH assay of S . palustris , antioxidant activity was higher in mature fertile (2.29 mg TE/mg GAE; TE: Trolox equivalent; GAE: gallic acid equivalent) and sterile (2.07 mg TE/mg GAE) fronds than in young fertile (1.99 mg TE/mg GAE) and sterile (1.80 mg TE/mg GAE) fronds. Moreover, significantly higher metal chelating activity was observed in young sterile fronds (101.22 µg EDTA/mg GAE; EDTA: ethylenediaminetetraacetic acid) and mature fertile fronds (176.37 µg EDTA/mg GAE) than in young fertile fronds (82.14 µg EDTA/mg GAE) and mature sterile fronds (16.54 µg EDTA/mg GAE) (Chai et al., 2012 2012 ). Methanol extracts from aerial plant parts of eleven different species of pteridophytes from Korea, namely A . pedatum , C . japonica , D . mariesii , C . fortunei , D . crassirhizoma , P . polyblepharum , P . lepidocolon , D . nipponensis , P . cretica , P. multifida , and P. nipponica displayed significant DPPH scavenging activity ranging from 0.05 to 1.66 mg/ml while the activity of rhizome extracts ranged only from 0.02 to 4.99 mg/ml. Using ABTS assays, Zakaria et al. ( 2011 2011 ) found 0.06 to 0.53 mg/ml of activity in methanol extracts from aerial plant parts and 0.03 to 1.24 mg/ml in rhizome extracts. In DPPH assays, methanol extracts from leaves of D . linearis exhibited higher antioxidant activity (20 µg/ml) (37.8%) than aqueous (29.3%) or chloroform extracts (13.3%). In D. linearis , superoxide scavenging activity was the highest in methanol extracts (84.2%), moderate in aqueous extracts (62.1%), and the lowest in chloroform extracts (23.2%) (Milan et al., 2013 2013 ). Methanol extracts of D. polypodiodes displayed only 5.87% antioxidant activity in DPPH assays (Kshirsagar and Upadhyay, 2009 2009 ). The ethyl acetate, butanol, and water fractions of B . orientale revealed significant free radical scavenging activity (IC 50 8.6–13.0 μg/ml) (Lai et al., 2010 2010 ). Significant DPPH scavenging (0.12 to 0.14 mg/ml) and ferric reducing power (FRP) (963 to 1417 mg GAE/100 g) were observed in methanol extracts from leaves of C. latebrosa , Cibotium barometz , D. quercifolia , B. orientale , and D. linearis . Adiantum raddianum and P . vittata displayed the highest activity in DPPH scavenging (0.27 and 0.29 mg/ml, respectively) and FRP (842 and 578 mg GAE/100 g, respectively) (Lai and Lim, 2011 2011 ). Anti-cancer, antidiabetic, and antiviral activity 2 The anti-cancer activity of ferns is listed in Table 3 3 , and their antidiabetic and antiviral activity in Table 4 4 . Both ethanol and water extracts of Hemionitis arifolia (Burm.) Moore (Fig. 2h 2h ) showed hypoglycaemic and antidiabetic properties in rats (Ajikumaran Nair et al., 2006 2006 ; Kumudhavalli and Jaykar, 2012 2012 ). The antihyperglycemic and analgesic activity of the leaves of Christella dentata , Adiantum philippense , and Angiopteris evecta has been studied by Paul et al. ( 2012 2012 ), Tanzin et al. ( 2013 2013 ), and Sultana et al. ( 2014 2014 ). Microsorum grossum has been used to cure liver cancer (Petard and Raau, 1972 1972 ; Defilpps et al., 1988 1988 ). Pteris polyphylla (Lee and Lin, 1988 1988 ) and A . evecta (Defilpps et al., 1988 1988 ) also show anti-cancer activity. In the Chinese traditional medicine system, “Gusuibu” has been used for its anti-cancer and anti-inflammation activity. It is prepared from the rhizomes of various ferns, namely D . fortunei (Kze.) J. Sm., P . coronans (Wall. Ex Mett.) Ching, D . divaricata BL., D. mariesii Moore ex-Bak, D. solida (Forst.) Sw., and H . griffithiana (Hk.) C. Chr. (Cui et al., 1990 1990 ; Cai et al., 2004 2004 ; ChPC, 2005 2005 ; Chang et al., 2007a 2007a ). M . quadrifolia has been used to treat human breast cancer, diabetes, and snake bites, and as an anti-inflammatory and diuretic (Uma and Pravin, 2013 2013 ). The bear paw fern, Phlebodium decumanum has been used for anti-cancer, ulcer, and high blood pressure treatments (Punzon et al., 2003 2003 ). The leaves and rhizomes of H . arifolia (Burm.) Moore and whole plants of Adiantum capillus-veneris L. were used by the tribes of the Valparai hills, Western Ghats, and Tamil Nadu (India) for their hypoglycaemic and anti-cancer activity (Santhosh et al., 2014 2014 ). Both leaves and rhizomes of A. capillus-veneris L. have been used in the preparation of herbal drugs for treating diabetes in India and Europe. This fern has diverse phytochemical constituents: adiantone, adiantoxide, astragalin, β-sitosterol, caffeic acids, caffeylgalactose, caffeylglucose, campesterol, carotenes, coumaric acids, coumarylglucoses, diplopterol, epoxyfilicane, fernadiene, fernene, filicanes, hopanone, hydroxy-adiantone, hydroxy-cinnamic acid, isoadiantone, isoquercetin, kaempferols, lutein, mutatoxanthin, naringin, neoxanthin, nicotiflorin, oleananes, populnin, procyanidin, prodelphinidin, quercetins, querciturone, quinic acid, rhodoxanthin, rutin, shikimic acid, violaxanthin, and zeaxanthin (Taylor, 2003 2003 ). MTT: 3-(4,5-dimethylthiazol-2-yl)-2,5-diphenyltetrazolium bromide No. Plant name Family name Methods used Source(s) 1 Microsorum grossum Polypodiaceae Traditional medicine Petard and Raau, 1972 1972 ; Defilpps et al., 1988 1988 2 Pteris polyphylla Pteridaceae In vitro antimutagenic activity, traditional Chinese medicine 2 Lee and Lin, 1988 1988 ; Wang and Zhang, 2008 2008 3 Pteris multifida Pteridaceae In vitro MTT method; antimutagenic activity against picrolonic acid-induced mutation 4 Angiopteris evecta Marattiaceae Traditional Chinese medicine Defilpps et al., 1988 1988 5 Drynaria fortunei Polypodiaceae Traditional Chinese medicine 3 Punzon et al., 2003 2003 ; Cai et al., 2004 2004 ; ChPC, 2005; Chang et al., 2007a 2007a 6 Pseudodrynaria coronans Polypodiaceae Traditional Chinese medicine 7 Phlebodium decumanum Polypodiaceae In vitro tumour necrosis inhibition 8 Davallia mariesii Davalliaceae Korea folk medicine 4 Cui et al., 1990 1990 ; Cai et al., 2004 2004 ; ChPC, 2005 9 Davallia divaricata Davalliaceae 3 Traditional Chinese medicine 10 Davallia solida Davalliaceae 11 Humata griffithiana Davalliaceae 12 Marsilea quadrifolia Marsileaceae In vitro cytotoxic activity on MCF-7 Uma and Pravin, 2013 2013 13 Hemionitis arifolia Pteridaceae 2 Traditional Indian medicine 2 Santhosh et al., 2014 2014 14 Adiantum capillus-veneris Pteridaceae 15 Blechnum orientale Blechnaceae In vitro MTT assay against human colonic adenocarcinoma HT-29, human colonic carcinoma HCT-116, human breast adenocarcinoma MCF-7, human leukemia K562 Lai and Lim, 2010 16 Polypodium nipponica Polypodiaceae 6 In vivo two-stage carcinogenesis test on mouse skin papillomas 8 Konoshima et al., 1996 1996 17 Polypodium formosanum Polypodiaceae 18 Polypodium vulgare Polypodiaceae 19 Polypodium fauriei Polypodiaceae 20 Polypodium virginianum Polypodiaceae 21 Dryopteris crassirizoma Dryopteridaceae 22 Adiantum monochlamys Pteridaceae 2 In vivo two-stage carcinogenesis test on mouse skin papillomas 23 Oleandra wallichii Oleandraceae 24 Cyrtomium fortunei Dryopteridaceae In vitro MTT assay on MGC-803, PC3, A375 tumor cells Yang et al., 2013 2013 25 Asplenium polyodon Aspleniaceae 5 Traditional Indian medicine 5 Singh, 1999 1999 ; Santhosh et al., 2014 2014 26 Asplenium nidus Aspleniaceae 27 Adiantum capillus-veneris Pteridaceae 28 Ophioglossum gramineum Ophioglossaceae 29 Hemionitis arifolia Pteridaceae 30 Pteridium aquilinum Dennstaedtiaceae Anticancer compounds identified Nwiloh et al., 2014 2014 31 Pityrogramma calomelanos Pteridaceae In vitro cytotoxicity against Dalton’s lymphoma ascites tumour cells and Ehrlich ascites tumour cells 3 Sukumaran and Kuttan, 1991 1991 ; Shin, 2010 2010 ; Zakaria et al., 2011 2011 ; Milan et al., 2013 2013 32 Drynaria quercifolia Polypodiaceae Brine Shrimp lethality bioassay 33 Dicranopteris linearis Gleicheniaceae In vitro MTT assay on MCT-7, HeLa, HT-29, HL-60, K-562, MDA-MB-231 34 Selaginella willdenowii Selaginellaceae In vitro cytotoxic activity on human cancer cell lines 10 Lee and Lin, 1988; Silva et al., 1995; Sun et al., 1997; Lee et al., 1999; Lin et al., 2000; Su et al., 2000; Chen et al., 2005b; Gao et al., 2007; Li J et al., 2014; Li S 2014; Elda et al., 2015 35 Selaginella lepidophyla Selaginellaceae In vitro MTT assay on human colon cancer cell line (HTB-38) 36 Selaginella labordei Selaginellaceae In vitro MTT assay on Bel-7402, HeLa, HT-29 37 Selaginella moellendorffii Selaginellaceae In vitro cytotoxic activity on human ovarian adenocarcinoma cells; in vitro MTT assay on cervical carcinoma (HeLa), foreskin fibroblast (FS-5), Bel-7402, HeLa, HT-29 38 Selaginella delicatula Selaginellaceae In vitro cytotoxic activity on Raji and Calu-1 tumor cell lines; in vitro MTT assay on Bel-7402, HeLa, HT-29 39 Selaginella tamariscina Selaginellaceae In vitro MTT assay; in vitro antiproliferative activity in leukemia cells, Bel-7402, HeLa, HT-29; in vivo single dose acute toxicity test 40 Selaginella doederleinii Selaginellaceae Antimutagenic activity against picrolonic acid-induced mutation; in vitro antiproliferative against human cancer cells 41 Selaginella uncinata Selaginellaceae 3 In vitro MTT assay on Bel-7402, HeLa, HT-29 42 Selaginella remotifolia Selaginellaceae 43 Selaginella pulvinata Selaginellaceae 44 Adiantum venustum Pteridaceae Ehrlich ascites carcinoma in animal models, in vivo animal study 3 Woerdenbag et al., 1996; Li et al., 1998, 1999; Viral et al., 2011 45 Pteris semipinnata Pteridaceae Human liver adenocarcinoma cell line (HePG II), human lung adenocarcinom a cell line (SPC-A-1), human gastric adenocarcinoma cell line (MGC-803), human nasopharyngeal carcinoma cells; in vitro MTT assay in HL-60 cells; in vivo anticancer activity 46 Pteris multifida Pteridaceae In vitro MTT assay in Ehrlich ascites 47 Pteris vittata Pteridaceae In vitro MTT assay on MCF-7 breast cancer cells 6 Chiu et al., 2009; Zhang et al., 2013; Tomsik, 2013; Kaur et al., 2014; Xia et al., 2014 48 Pteridium aquilinum Dennstaedtiaceae In vitro MTT assay 49 Davallia cylindrica Davalliaceae 2 In vitro MTT assay in A549 cells 50 Stenoloma chusanum Lindsaeaceae 51 Selaginella frondosa Selaginellaceae In vitro MTT assay in A549 cells 52 Thelypteris torresiana Thelypteridaceae In vitro MTT assay in lung cancer cell line H1299 53 Selaginella ciliaris Selaginellaceae 3 Potato disc-induced method 3 Sarker et al., 2011 54 Marsilea minuta Marsileaceae 55 Thelypteris prolifera Thelypteridaceae 56 Trichomanes reniforme Hymenophyllaceae In vivo growth inhibitory activity against ascitic fibrosarcoma in Swiss mice;in vitro cytotoxicity Guha et al., 1996 Medicinal use No. Plant name Family name Methods used Source(s) 30 Antiviral activity 1 Hemionitis arifolia Pteridaceae In vivo glucose tolerance test and alloxan-induced diabetic rats; in vivo glucose tolerance test in streptozotocin-induced diabetic rats; traditional Indian medicine Ajikumaran Nair et al., 2006 2006 ; Kumudhavalli and Jaykar, 2012 2012 ; Santhosh et al., 2014 2014 2 Christella dentata Thelypteridaceae In vivo glucose tolerance test 3 Paul et al., 2012 2012 ; Tanzin et al., 2013 2013 ; Sultana et al., 2014 2014 3 Adiantum philippense Pteridaceae In vivo study in alloxan-induced diabetic rats 4 Angiopteris evecta Marattiaceae In vivo glucose tolerance test 5 Adiantum capillus-veneris Pteridaceae Traditional Indian medicine; traditional medicine in Amazon; in vivo glucose tolerance test Jain and Sharma, 1967 1967 ; Taylor, 2003 2003 ; Santhosh et al., 2014 2014 ; Neef et al., 1995 1995 6 Angiopteris evecta Marattiaceae In vivo glucose tolerance test 2 Miao et al., 1996 1996 ; Nguyen, 2005 2005 7 Selaginella tamarascina Selaginellaceae In vivo study in alloxan-induced diabetic rats 8 Asplenium polyodon Aspleniaceae 5 Traditional Indian medicine 6 Singh, 1999 1999 ; Chand Basha et al., 2013 2013 ; Santhosh et al., 2014 2014 9 Asplenium nidus Aspleniaceae 10 Adiantum capillus-veneris Pteridaceae 11 Ophioglossum gramineum Ophioglossaceae 12 Hemionitis arifolia Pteridaceae 13 Actiniopteris radiata Pteridaceae In vitro chromogenic DNSA method 14 Blechnum orientale Blechnaceae 8 In vitro antiviral activity against hepatitis B virus (HBV), hepatitis C virus (HCV), human immunodeficiency virus (HIV), and murine leukemia virus 8 Tsai and Hwang, 1999 1999 15 Woodwardia orientalis Blechnaceae 16 Woodwardia unigemmata Blechnaceae 17 Athyrium acrostichoides Athyriaceae 18 Sphaeropteris lepifera Cyatheaceae 19 Cyrtomium fortunei Dryopteridaceae 20 Dryopteris crassirhizoma Dryopteridaceae 21 Osmunda japonica Osmundaceae 22 Lygodium flexuosum Lygodiaceae Traditional Indian medicine Dhiman, 1998 1998 23 Lygodium japonicum Lygodiaceae In vitro antiviral activity against herpes simplex virus, Sindbis virus, and polio virus Taylor et al., 1996 1996 24 Asplenium adiantum-negrum Aspleniaceae Traditional Indian medicine Vasudeva, 1999 1999 25 Microsorum membranifolium Polypodiaceae Traditional Fijian medicine Cambie and Ash, 1994 1994 26 Pyrrosia lingua Polypodiaceae In vitro antiviral activity against Herpes simplex virus Zheng, 1990 1990 27 Trichomanes reniforme Hymenophyllaceae In vitro antiviral activity against HIV Guha et al., 1996 1996 28 Pteris glycyrrhiza Pteridaceae 3 In vitro antiviral activity against bovine herpesvirus type McCutcheon et al., 1995 1995 29 Polypodium vulgare Pteridaceae 2 Husson et al., 1986 1986 30 Polypodium aureum Pteridaceae 6 2 Antiviral activity 31 Athyrium niponicum Athyriaceae In vitro antiviral activity 32 Selaginella uncinata Selaginellaceae In vitro antiviral activity against respiratory syncytial virus, parainfluenza type 3 virus 6 21 Anti-inflammatory activity 33 Microsorum grossum Polypodiaceae Traditional herbal medicine 34 Drynaria quercifolia Polypodiaceae Traditional Indian medicine 35 Microsorum scolopendria Polypodiaceae Traditional herbal medicine in Tonga 36 Polypodium leucotomos Polypodiaceae Traditional herbal medicine in Bolivia 37 Ophioglossum vulgatum Ophioglossaceae 3 Traditional Indian medicine 38 Lygodium flexuosum Lygodiaceae 39 Adiantum venustum Pteridaceae 40 Adiantum caudatum Pteridaceae Traditional medicine in Malay Peninsula 41 Cheilanthes farinosa Pteridaceae In vivo anti-inflammatory activity 42 Pteris ensiformis Pteridaceae In vitro anti-inflammatory activity 43 Christella parasitica Thelypteridaceae Traditional Indian medicine 44 Huperzia serrata Huperziaceae Traditional Chinese medicine 45 Selaginella tamarascina Selaginellaceae In vitro anti-inflammatory activity 46 Selaginella bryopteris Selaginellaceae Traditional Indian medicine 47 Dryopteris sp Dryopteridaceae 2 In vivo Carrageenan induced paw oedema method 48 Blechnum occidentale Blechnaceae 49 Alsophila gigantea Cyatheaceae Traditional Indian medicine 50 Cyathea gigantea Cyatheaceae In vivo anti-inflammatory activity 51 Phlebodium decumanum Polypodiaceae In vitro anti-inflammatory activity 52 Phyllitis scolopendrium Aspleniaceae Traditional herbal medicine 53 Dicranopteris linearis Gleicheniaceae In vitro anti-inflammatory activity 6 11 Wound healing activity 54 Selaginella delicatula Selaginellaceae 2 Traditional Indian medicine 55 Dicranopteris linearis Gleicheniaceae 56 Davallia solida Davalliaceae Traditional medicine in Hawaii 57 Polystichum pungens Dryopteridaceae Traditional medicine in South Africa 58 Angiopteris evecta Marattiaceae Traditional medicine in Fijian 59 Nephrolepis cordifolia Nephrolepidaceae 3 Traditional Indian medicine 60 Ophioglossum reticulatum Ophioglossaceae 61 Thelypteris arida Thelypteridaceae 62 Phyllitis scolopendrium Aspleniaceae In vivo method 63 Pityrogramma calomelanos Pteridaceae Traditional medicine 64 Polypodium ssp. Polypodiaceae In vitro assay using platelet activating factor and leukotriene B(4) Pterosins, terpenoids, sterides, flavones, glucosides, aromatics, and pyrone compounds derived from C . barometz have antitumour properties (Wu et al., 2007 2007 ; Xu et al., 2012 2012 ). Plants in the polyploid genus Pteris contain many chemical compounds like apigenin-7- O -β- D -glucopyranosyl-4'- O -α- L -rhamnopyranoside, luteolin-7- O -β- D -glucopyranoside, apigenin-7- O -β- D - glucopyranoside, apigenin, luteolin, naringenin-7- O -β- D -neohesperidoside, apigenin-7- O -β- D -neohespedoside, apigenin-4'- O -α- L -rhamnopyranoside, (2S,3S)-pterosin C, isovanillic acid, ferulic acid, 2β,16α-dihydroxy-ent-kaurane 2- O -β- D -glucoside, 2β,6β,15α-trihydroxy-ent-kaur-16-ene, 9-hydroxy-15-oxo-ent-kaur-16-en-19-oic acid, 19β- D -glucoside, 2β,14β,15α,16α,17-pentahydroxy-ent-kaurane, and 9-hydroxy-ent-kaur-16-en-19-oic acid (Gong et al., 2007 2007 ; Ouyang et al., 2008 2008 ). Extracts of P. multifida have anti-cancer activity (Wang and Zhang, 2008 2008 ). Ethyl acetate, butanol, and water extracts of B . orientale (Linn.) show notable anti-cancer properties (Lai et al., 2010 2010 ). The antithiamin substances in brackens, such as astragalin, isoquercitrin, rutin, caffeic acid, and tannic acid have been identified as valuable natural constituents because of their antitumour or antioxidant properties (Kweon, 1986 1986 ; Cai et al., 2004 2004 ; Katsube et al., 2006 2006 ). The ferns P . nipponica , Polypodium formosanum , Polypodium vulgare , Polypodium fauriei , Polypodium virginianum , Dryopteris crassirizoma , Adiantum monochlamys , and Oleandra wallichii have been shown to have anti-cancer activity (Konoshima et al., 1996 1996 ). A . evecta and Selaginella tamariscina have been shown to have significant antidiabetic activity (Miao et al., 1996 1996 ; Nguyen, 2005 2005 ). According to Singh ( 1999 1999 ) and Lee et al. ( 2003 2003 ), Asplenium polyodon , Asplenium nidus , A . capillus-veneris , Ophioglossum gramineum , and H . arifolia have anti-cancer, antidiabetic, and antiviral activity. Chloroform extracts of Actiniopteris radiata (Linn.) have significant antidiabetic activity as shown by the chromogenic DNSA (dinitrosalicylic) colorimetric method, and ethanolic extracts show strong in vitro antidiabetic properties when compared to n -hexane (Chand Basha et al., 2013 2013 ). Both ethyl acetate and butanol extracts of the rhizome of C . fortunei (J.) Smith (Fig. 3a 3a ) exhibit anti-tumour activity (Yang et al., 2013 2013 ). The phlorophenone derivatives in Dryopteris sp. (Kapadia et al., 1996 1996 ) and A . capillus-veneris (Fig. 3b 3b ) that have anti-cancer activity have also been used as antidiabetic drugs in India and Europe (Jain and Sharma, 1967 1967 ; Neef et al., 1995 1995 ). β-Caryophyllene, linalool, geranial and γ-terpinene have also been isolated from the angiosperms Citrus and Croton flavens , revealing antitumour and antioxidant properties (Choi et al., 2000 2000 ; Sylvestre et al., 2006 2006 ). Moreover, anti-cancer compounds were detected in the essential oils of P . aquilinum L. Kuhn (Fig. 2a 2a ) (Nwiloh et al., 2014 2014 ). Methanol extracts of Pityrogramma calomelanos , D . quercifolia , and D . linearis exhibit antitumour and cytotoxic activity (Sukumaran and Kuttan, 1991 1991 ; Shin and Lee, 2010 2010 ; Zakaria et al., 2011 2011 ; Milan et al., 2013 2013 ). Extracts of Adiantum venustum also have anti-cancer activity, as demonstrated by Viral et al. ( 2011 2011 ). The lycophytes, Selaginella willdenowii (Fig. 3d 3d ), Selaginella lepidophyla , Selaginella labordei , Selaginella moellendorffii , Selaginella delicatula , S. tamariscina , and Sellaginella doederleinii (Fig. 3e 3e ) were shown to have cytotoxic and antimutagenic effects due to the presence of bioflavonoids (Lee and Lin, 1988 1988 ; Silva et al., 1995 1995 ; Sun et al., 1997 1997 ; Lee et al., 1999 1999 ; Lin et al., 2000 2000 ; Su et al., 2000 2000 ; Chen JJ et al., 2005 2005 ; Gayathri et al., 2005 2005 ; Woo et al., 2005 2005 ; Gao et al., 2007 2007 ; Shi et al., 2008 2008 ; Tan et al., 2009 2009 ), while Pteris semipinnata (Fig. 4c 4c ) and P. multifida have cytotoxic effects due to the presence of diterpenes (Woerdenbag et al., 1996 1996 ; Li et al., 1998 1998 , 1999 1999 ). Several authors have studied the anti-cancer potentials of ferns and lycopods, namely P . aquilinum , Davallia cylindrical , Stenoloma chusanum , Selaginella frondosa , Thelypteris torresiana , P . vittata , Selaginella ciliaris , Marsilea minuta , and Thelypteris prolifera (Chiu et al., 2009 2009 ; Sarker et al., 2011 2011 ; Tomsik, 2013 2013 ; Zhang et al., 2013 2013 ; Kaur et al., 2014 2014 ; Xia et al., 2014 2014 ). Note: for interpretation of the references to color in this figure legend, the reader is referred to the web version of this article According to Tsai and Hwang ( 1999 1999 ), the hepatitis B virus (HBV) is a prominent infectious disease agent worldwide, especially in Asia and Africa, causing liver cirrhosis and cancer. Blechni rhizome, known for its antiviral (HBV) activity, is prepared from the roots and stems of various ferns including B . orientale , D . crassirhizoma , O . japonica , Woodwardia orientalis (Fig. 4d 4d ), Woodwardia unigemmata (Fig. 5b 5b ), Athyrium acrostichoides , Sphaeropteris lepifera , and C . fortunei . Likewise, an herbal mixture of Woodwardia species with some other flowering plants has shown anti-HIV-1 (human immunodeficiency virus) activity. Lygodium flexuosum (Dhiman, 1998 1998 ) and Asplenium adiantum-nigrum (Vasudeva, 1999 1999 ) have shown significant effects against viral disease and jaundice, while Microsorum membranifolium has shown activity against children’s influenza (Cambie and Ash, 1994 1994 ). The antiviral and anti-tumor activity shown by Trichomanes reniforme is due to the presence of mangiferin, which can kill cancer cells and has anti-HIV effects (Guha et al., 1996 1996 ). The antiviral activity of Pteris glycyrrhiza (McCutcheon et al., 1995 1995 ), P. vulgare , and Polypodium aureum (Husson et al., 1986 1986 ) has been documented. A new antiviral compound, woodorien, was derived and identified by Xu et al. ( 1993 1993 ). A . niponicum is reported to have anti-HIV properties (Mizushina et al., 1998 1998 ), while Taylor et al. ( 1996 1996 ) have shown that L . japonicum has antiviral effects against the Sindbis virus. Zheng ( 1990 1990 ) revealed the antiviral activity of P . lingua (Fig. 3g 3g ) against the herpes simplex virus. Selaginella uncinata (Fig. 5c 5c ) displays antiviral activity against the respiratory syncytial virus (RSV) and parainfluenza type 3 virus (PIV3) (Ma et al., 2003 2003 ). Note: for interpretation of the references to color in this figure legend, the reader is referred to the web version of this article Anti-inflammatory activity 2 In general, free radicals participate in numerous natural processes that are harmful to lipids, proteins, membranes, and nucleic acids, thereby contributing to a variety of diseases (Hadi et al., 2000 2000 ; Lee et al., 2005 2005 ; Campos et al., 2006 2006 ). Reactive oxygen species (ROS) contribute to both normal cell function and pathological ailments such as atherosclerosis, inflammatory injury, cancer, and cardiovascular disease. Inflammation is a typical defensive response to cellular damage produced by physical distress (cuts, burns, or bruises), toxic chemicals, microbial agents, or autoimmune disease. Unfortunately, drugs currently available to treat pain and inflammation have numerous harmful side effects and low efficiency. Natural products are often easily available, with high clinical effectiveness and few side effects. The World Health Organization (WHO) estimates that 80% of the global population still depends on ethnomedicinal practices for treating their illnesses (Gurib-Fakim, 2006 2006 ). In inflammatory disorders, extreme stimulation of phagocytes and the production of O 2− (oxygen radicals), OH − (hydroxide ions), and H 2 O 2 (Gillham et al., 1997 1997 ) are severely harmful to tissues, by their significant direct or indirect oxidizing action. Hydrogen peroxide and OH – radicals formed from O 2− cause lipid peroxidation resulting in membrane destruction. In addition, tissue damage provokes an inflammatory response involving the production of mediators and chemotactic factors (Lewis, 1989 1989 ). The free radical scavenging activity of plant products is primarily due to various bioactive compounds like phenolics, flavonoids, phenolic diterpenes, and tannins (Lee et al., 2004 2004 ). Currently, steroidal and nonsteroidal anti-inflammatory drugs (NSAIDs) used to treat inflammatory disease frequently produce severely harmful effects (Vonkeman and van de Laar, 2010 2010 ; Vijayalakshmi et al., 2011 2011 ). Thus, there is abundant motivation to find new drugs with anti-inflammatory properties and fewer side effects, especially from plants used in folk medicine. Moreover, plant-derived natural products such as alkaloids, flavonoids, tannins, and terpenes have attracted significant attention recently due to their varied medicinal actions, including anti-inflammatory, analgesic, and antioxidant properties (Pandurangan et al., 2008 2008 ). Traditional healers have defined the healing efficiencies of native medicinal plants for various diseases. Because bioproducts are a source of synthetic and traditional herbal medicines, which can potentially be used by modern health care systems, scientists have been urged to study these plants to identify their medicinal properties (Vimala et al., 2012 2012 ). Pteridophytes (the ferns and their allies) have an extensive geologic history as pioneer plants that have occupied various parts of our globe for millions of years. This group comprises a huge group of seedless vascular plants and occupies a significant place in primary health care because of their cost effectiveness. Economic crises in developing countries mean that different fern species are presently used for various ailments (Ho et al., 2011 2011 ). On the basis of antioxidant activity, ferns exhibit bioactivity such as anti-microbial, anti-viral, anti-inflammatory, anti-tussive, and anti-tumor activity (Chang et al., 2011 2011 ). The Malasar tribes in the Valparai hills (Western Ghats, India) use both S . delicatula (Fig. 5a 5a ) and D . linearis for their wound healing activity (Santhosh et al., 2014 2014 ). Table 4 4 shows the anti-inflammatory and wound healing activity of pteridophytes. In many studies, anti-inflammatory activity has been reported for many species of medicinal ferns, including M . grossum (Petard and Raau, 1972 1972 ; Whistler, 1992 1992 ), D . quercifolia (Dixit and Vohra, 1984 1984 ), A . aureum (Defilpps et al., 1988 1988 ), Selaginella bryopteris (Dhiman, 1998 1998 ), Ophioglossum vulgatum , L . flexuosum , A . venustum (Vasudeva, 1999 1999 ), Adiantum caudatum (Burkill, 1996 1996 ), Christella parasitica (Gogoi, 2002 2002 ), Huperzia serrata (Fig. 5d 5d ) (Zangara, 2003 2003 ), S . tamariscina (Woo et al., 2005 2005 ), P . ensiformis (Fig. 4f 4f ) (Wu et al., 2005 2005 ), P. multifida (Lee and Lin, 1988 1988 ), Dryopteris sp. (Otsuka et al., 1972 1972 ), Cheilanthes farinosa (Yonathan et al., 2006 2006 ), Cyathea gigantea (Fig. 3c 3c ) (Benjamin and Manickam, 2007 2007 ; Madhukiran and Ganga Rao, 2011 2011 ), Blechnum occidentale (Nonato et al., 2009 2009 ), D . mariesii (Chang et al., 2007b 2007b ), Microsorum scolopendria (Bloomfield, 2002 2002 ), Phyllitis scolopendrium (Bonet and Valles, 2007 2007 ), and P . leucotomos (Lucca, 1992 1992 ). In addition, wound healing activity has been reported for many species of ferns such as D . solida (Whistler, 1992 1992 ), Polystichum pungens (Grierson and Afolayan, 1999 1999 ), A . evecta (Cambie and Ash, 1994 1994 ), Nephrolepis cordifolia (Fig. 5e 5e ), Ophioglossum reticulatum , Thelypteris arida (Upreti et al., 2009 2009 ), P . scolopendrium (Oniga et al., 2004 2004 ), P . calomelanos (Fig. 5f 5f ) (de Feo, 2003 2003 ), Polypodium spp. (Liu et al., 1998 1998 ), and P . decumanum (Punzon et al., 2003 2003 ). Antimicrobial activity 2 There are many antibiotics currently available for the treatment of infection, including chloramphenicol, tetracycline, erythromycin, penicillin G, ampicillin, cephalosporin, ciprofloxacin, kanamycin, gentamicin, neomycin, amoxycillin, nystatin, amphotericin-B, and ketaconazole (Kaufman, 2000 2000 ). Although there are numerous antibiotics for healing bacterial and fungal infection, they are not always reliable against pathogenic organisms (Gearhart, 1994 1994 ). Their use is restricted by several factors such as low potency, poor solubility, and drug toxicity (Fromtling and Rahway, 1987 1987 ; Portillo et al., 2001 2001 ). The development of resistance in pathogenic bacteria and fungi against most of the conventionally available drugs has been reported (Cuenca-Estrella et al., 2000 2000 ). Confounding the issue further is the indiscriminate use of antibiotics, which leads to the loss of their effectiveness. Many of the pathogens, especially bacteria and fungi, have developed substantial resistance to the antibiotics (Darokar et al., 1998 1998 ; Jones, 1998 1998 ; Austin et al., 1999 1999 ). Moreover, the widespread use of antibiotics has led to the decimation of sensitive organisms from the population with the consequence that there has been an increase in the number of resistant microorganisms. This situation has forced scientists to search for new bioactive compounds in traditional complementary medicines derived from plants (Kumar et al., 2006 2006 ). Plant extracts have been widely used since ancient times for treating human illnesses. Numerous studies of the antimicrobial activity of different natural products have been reported (Bhattacharjee et al., 2006 2006 ; Parekh and Chanda, 2006 2006 , 2007 2007 ; Vaghasiya et al., 2008 2008 ). Although the medicinal value of pteridophytes has been known to traditional cultures for more than 2000 years, they are used only on a small scale in modern chemotherapy. Studies of the bioactivity of pteridophytes are still in their early years compared with those of angiosperms. The Greek botanist Theophrastus (ca. 372–287 BC) wrote about the medicinal value of pteridophytes. Dioscorides (ca. 50 AD) in his de Materia Medica also referred to a number of ferns, including P . aquilinum and Dryopteris filix-mas , as having medicinal values. In ancient Indian medicine systems, several ferns were used to cure a number of human ailments. Sushruta (ca. 100 AD) and Charaka (ca. 100 AD) recommended the medicinal use of some ferns in their samhitas. Several ferns have been used by Unani physicians in India and Western Asia (Banerjee and Sen, 1980 1980 ). The antimicrobial activity of pteridophytes is summarized in Table 5 5 . Several studies have been carried out to explore the antimicrobial activity of ferns such as Nephrolepis acuminata (Jimenez et al., 1979 1979 ), Davallia sodila , Lygodium reticulatum (Cambie and Ash, 1994 1994 ), Marattia fraxinea (de Boer et al., 2005 2005 ), Sphenomeris chinensis (Sengupta et al., 2002 2002 ), A . caudatum , Adiantum peruvianum , A. venustum , Adiantum incisum , Adiantum latifolium , and Ampelopteris prolifera (Banerjee and Sen, 1980 1980 ; Lakshmi et al., 2006 2006 ; Lakshmi and Pullaiah, 2006 2006 ; Singh et al., 2008a 2008a ), P . aquilinum (Francisco and Driver, 1984 1984 ), Nephrolepis sp. (Basile et al., 1997 1997 ), Adiantum lunulatum (Reddy et al., 2001 2001 ), E . arvense (Joksic et al., 2003 2003 ; Radulovic et al., 2006 2006 ), S . tamariscina (Woo et al., 2005 2005 ), A . capillus-veneris (Guha et al., 2004 2004 , 2005 2005 ; Besharat et al., 2008 2008 ), Athyrium pectinatum (Parihar et al., 2006 2006 ), P . vittata (Singh et al., 2008b 2008b ), P. multifida (Hu et al., 2008 2008 ; Hum et al., 2008 2008 ), Mecodium exsertum (Maridass, 2009 2009 ), Selaginella involvens (Fig. 5g 5g ), S. inaequalifolia (Haripriya et al., 2010 2010 ), S. pallescens (Rojas et al., 1999 1999 ), Asplenium scolopendrium , Cystopteris fragilis , P . vulgare (Soare et al., 2012b 2012b ), A . caudatum , A . evecta , Pteris confusa , P. argyraea , Lygodium microphyllum (Gracelin et al., 2012 2012 ), Pteris biaurita (Dalli et al., 2007 2007 ; de Britto et al., 2012 2012 ), D . crassirhizoma (Lee et al., 2009 2009 ), and various species (Maruzzella, 1961 1961 ; McCutcheon et al., 1995 1995 ). Furthermore, several studies (May, 1978 1978 ; Dixit and Vohra, 1984 1984 ; Dixit, 1992 1992 ; Verma and Singh, 1995 1995 ; Manandhar, 1996 1996 ; Das, 1997 1997 ; Singh, 1999 1999 ; Benjamin and Manickam, 2007 2007 ) have reported the antiviral, antibacterial, and antifungal activity of Equisetum ramosissimum (Fig. 4a 4a ), A . aureum , D . quercifolia , Psilotum nudum (Fig. 5h 5h ), Parahemionitis arifolia , Helminthostachys zeylanica , O . gramineum , Tectaria caodunata , S . involvens , S. delicatula , Hypodematium crenatum , Leucostegia immersa , and Solvinia molusta . Singh ( 1999 1999 ) documented antibacterial and antifungal activity of Botrychum lanuginosum , Dryopteris cochleata , N . cordifolia , Polystichum molluscens , Polystichum squarrosum , Salvinia radicata , Sphaerostephanos unitus , S . palustris , P . cretica , and P . vittata . Although several studies have reported the antimicrobial activity of ferns, in-depth studies to estimate minimum inhibitory concentrations (MIC) and minimum bactericidal concentrations (MBC) have not yet been carried out for several ferns. No. Plant name Family name Methods used Source(s) 1 Nephrolepis acuminata Nephrolepidaceae In vitro bioassay Jimenez et al., 1979 1979 2 Davallia sodila Davalliaceae 2 Traditional Fijian medicine 2 Cambie and Ash, 1994 1994 3 Lygodium reticulatum Lygodiaceae 4 Marattia fraxinea Marattiaceae 16 In vitro antimicrobial assay de Boer et al., 2005 2005 5 Sphenomeris chinensis Lindsaeaceae Sengupta et al., 2002 2002 6 Adiantum caudatum Pteridaceae 6 Banerjee and Sen, 1980 1980 ; Lakshmi and Pullaiah, 2006 2006 ; Lakshmi et al., 2006 2006 ; Singh et al., 2008a 2008a 7 Adiantum peruvianum Pteridaceae 8 Adiantum venustum Pteridaceae 9 Adiantum incisum Pteridaceae 10 Adiantum latifolium Pteridaceae 11 Ampelopteris prolifera Thelypteridaceae 12 Pteridium aquilinum Dennstaedtiaceae Francisco and Driver, 1984 1984 13 Nephrolepis sp. Nephrolepidaceae Basile et al., 1997 1997 14 Equisetum arvense Equisetaceae Joksic et al., 2003 2003 ; Radulovic et al., 2006 2006 15 Athyrium pectinatum Athyriaceae Parihar et al., 2006 2006 16 Adiantum lunulatum Pteridaceae Reddy et al., 2001 2001 17 Adiantum capillus-veneris Pteridaceae Guha et al., 2004 2004 , 2005 2005 ; Besharat et al., 2008 2008 18 Pteris vittata Pteridaceae Singh et al., 2008b 2008b 19 Pteris multifida Pteridaceae Hu et al., 2008 2008 ; Hum et al., 2008 2008 20 Microsorum grossum Polypodiaceae Traditional Polynesian medicine Whistler, 1992 1992 21 Mecodium exsertum Hymenophyllaceae 14 In vitro antimicrobial assay Maridass, 2009 2009 22 Selaginella involvens Selginellaceae 3 Rojas et al., 1999 1999 ; Haripriya et al., 2010 2010 23 Selaginella inaequalifolia Selginellaceae 24 Selaginella pallescens Selginellaceae 25 Asplenium scolopendrium Aspleniaceae 3 Soare et al., 2012b 2012b 26 Cystopteris fragilis Cystopteridaceae 27 Polypodium vulgare Polypodiaceae 28 Angiopteris evecta Marattiaceae 5 Gracelin et al., 2012 2012 29 Adiantum caudatum Pteridaceae 30 Pteris confusa Pteridaceae 31 Pteris argyraea Pteridaceae 32 Lygodium microphyllum Lygodiaceae 33 Pteris biaurita Pteridaceae Dalli et al., 2007 2007 ; de Britto et al., 2012 2012 34 Dryopteris crassirizoma Dryopteridaceae Lee et al., 2009 2009 35 Equisetum ramosissimum Equisetaceae 13 Traditional Indian medicines 13 May, 1978 1978 ; Dixit and Vohra, 1984 1984 ; Dixit, 1992 1992 ; Verma and Singh, 1995 1995 ; Manandhar, 1996 1996 ; Das, 1997 1997 ; Singh, 1999 1999 ; Benjamin and Manickam, 2007 2007 36 Drynaria quercifolia Polypodiaceae 37 Psilotum nidum Psilotaceae 38 Parahemionitis arifolia Pteridaceae 39 Acrostichum aureum Pteridaceae 40 Helminthostachys zeylanica Ophioglossaceae 41 Ophioglossum gramineum Ophioglossaceae 42 Tectaria caodunata Tectariaceae 43 Selaginella involvens Selaginellaceae 44 Selaginella delicatula Selaginellaceae 45 Hypodematium crenatum Woodsiaceae 46 Leucostegia immersa Davalliaceae 47 Solvinia molusta Salviniaceae 48 Marsilea quadrifolia Marsileaceae Brine shrimp lethality bioassay Ripa et al., 2009 2009 Activity against Alzheimer’s disease and brain illness 2 Frequently, alkaloids act as stimulators of the nervous system, and occasionally as toxins. Cocaine (has an anaesthetic effect), atropine (affects motor nerves), and curare (used by South American natives to cause paralysis of prey), are all alkaloids (Kretovich, 1966 1966 ). Even though several alkaloids are present in angiosperms, some specific groups of alkaloids, namely lycopodine and huperzine, are synthesized by the lower groups of plants, like club mosses. Some lycopodium alkaloids naturally present in Lycopodium sp. and other pteridophytes have some pharmacological activity. α-Onocerin and lycoperine A show acetylcholinesterase inhibition activity (Zhang et al., 2002 2002 ; Hirasawa et al., 2006 2006 ). Huperzine A obtained from Huperzia species and other members of the Lycopodiaceae, has been shown to improve memory in animals. Huperzia quadrifariata and Huperzia reflexa were also found to cure Alzheimer’s disease (Ma and Gang, 2004 2004 ; Ma et al., 2007 2007 ; Singh and Singh, 2010 2010 ; Konrath et al., 2012 2012 ; Yang et al., 2012 2012 ). Amentoflavone and ginkgetin, flavonoids found in Selaginella , display neurodefensive activity against cytotoxic stress, which suggests their promising usage to cure neurodegenerative diseases such as stroke and Alzheimer’s (Kang et al., 2004 2004 ). The neuromodulatory tendency of S. delicatula using in vivo models of chemically induced neurodegenerative diseases has been studied in rodent systems and Drosophila . Extracts of S. delicatula showed neuroprotective efficacy against Parkinson’s disease, Huntington’s disease, reduced glutathione levels, and acetyl cholinesterase inhibitory activity (Chandran and Muralidhara, 2014 2014 ). The pteridophyte Drynaria is also used in traditional medicine in China to treat Alzheimer’s disease. According to Yang ( 2005 2005 ), 120 g of Drynaria (in decoction) produced significant improvement in Alzheimer’s disease patients. Likewise, Nephrolepis auriculata , S . bryopteris , H. zeylanica , and Lycopodium were used to treat brain fever, headache, and unconsciousness, to improve vitality, and as a brain tonic and memory enhancer (Bai, 1993 1993 ; Samant et al., 1998 1998 ; Vasudeva, 1999 1999 ; Kholia and Punetha, 2005 2005 ). Traditional uses of some pteridophytes 1 In India, the Gond communities of Badkachhar, Bariaam and Dapka ingest the juice of whole fresh plants to cure cough and diabetes mellitus. Leaves of A . philippense are used to cure fever, cough, asthma, leprosy, and hair loss, while whole plants of B . orientale are used in anathematic and typhoid treatments. Fronds of B . orientale ground in cow’s milk are used in the treatment of asthma, as antibacterials and anthelmintics, and to improve fertility in women. The whole plant of H . zeylanica is considered an intoxicant, anodyne, and is also used as an aphrodisiac. L . flexuosum (Fig. 4e 4e ) is used to cure skin diseases, rheumatism, sprains, scabies, eczema, cut wounds, and rheumatism, and as an expectorant. A paste made from the leaves of N . cordifolia is applied to wounds to stop bleeding. Whole plants of S . bryopteris are used as diuretics, and to treat gonorrhoea (Singh and Upadhyay, 2012 2012 ). A . capillus-veneris is used for antifungal growth and to remove dandruff, and the juice of leaves is drunk to protect against infertility. In addition, A . caudatum is used to cure jaundice, scabies, abdominal pain, and constipation. The young fronds of A . incisum are used to treat malaria and bronchial diseases, while leaves and rhizomes of A. venustum are thought to cure diabetes, and are used to treat liver problems and circulatory disorders. Leaves of Asplenium bulbiferum (Fig. 4b 4b ) are applied externally to hemorrhoids, and fronds are taken for liver problems. Decoctions from the rhizomes of P . aquilinum are drunk as a herbal health tea, and the rhizomes of W . unigemmata are used to cure skin diseases. Leaf juices are used as a tea or bath for curing infertility in women, for abdominal pain, constipation and sore throats (Nwosu, 2002 2002 ). A paste of leaves and rhizomes mixed with lukewarm milk for 30 d is used as an effective tonic. A paste of P . vittata was applied twice to glandular swellings for 15 d (Sharma, 2002 2002 ). M . grossum is commonly used in Tahiti (Whistler, 1992 1992 ), H . serrata in China (Zangara, 2003 2003 ), and S . bryopteris in India (Dhiman, 1998 1998 ) for their anti-inflammatory activity. A decoction of roots and fronds of D . quercifolia is used as a tonic for the bowels, and aqueous extracts of the rhizomes are applied to wounds and cuts (Anonymous, 1952 1952 , 1986 1986 ; Singh, 1973 1973 ; Islam, 1983 1983 ; Jain, 1991 1991 ). Dried rhizomes of P . aquilinum mixed with milk are used to relieve diabetic disorders, and tender fronds are used as vegetables. Green fronds are used as fodder, make a good soil binder, and are also used in the preparation of manure (Anonymous, 1972 1972 , 1986 1986 ; Islam, 1983 1983 ; Jain, 1991 1991 ). A paste made from the rhizomes of O . vulgatum has been externally applied to wounds, dropped in sore eyes as a detergent, and made into an antiseptic. Infusions can be taken internally for vulnerary or to treat bleeding of the nose or mouth, and a decoction has been drunk to treat heart troubles (Anonymous, 1966 1966 , 1986 1986 ; Islam, 1983 1983 ). In Malaya, a rhizome paste of A . aureum is applied to wounds and boils, and in the Philippines, leaves of A . caudatum are used externally as remedies for skin disease and internally for diabetes. Rhizomes of C . barometz in the Philippines are used to treat wounds and ulcers. Roots and leaves of Lygodium circinnatum are applied to wounds in the Dutch East Indies (Quisumbing, 1951 1951 ). Tribal people of the Palani hills from South India use A . incisum , C . gigantea , and C. nilgirensis to treat diabetes, while N . cordifolia , H . arifolia , and Microsorum punctatum are applied to cuts and wounds, or used as anti-inflammatories (Sathiyaraj et al., 2015 2015 ). In the Similipal Biosphere Reserve of Orissa (India), A . evecta , A . caudatum , B . orientale , Ceratopteris thalictroides , M . punctatum , P . biaurita , and P. cretica are used to cure cuts, wounds, and inflammation (Rout et al., 2009 2009 ). Mechanisms of action of phytochemicals 1 In this review we have documented most of the pteridophytes evaluated for phytochemical composition and bioactivity. Tables 1 1 – 5 5 summarize the presence of phytochemicals and their medicinal uses in pteridophytes. Although there are more than 12 000 species of pteridophytes distributed all over the world, few have been evaluated for their phytochemicals and their pharmacological properties. Most studies have reported only in vitro bioactivity, which suggests to investigate the pteridophytes could exert in vivo bioactivity. In general, phenolic compounds exist in both edible and non-edible plants and have proven bioactive. Phenolics are present in diverse groups of plants and show antimutagenic, anti-cancer, and antioxidant activity, which is beneficial to human life (Li et al., 2006 2006 ). Due to synergistic effects, mixtures of bioactive compounds such as phenols, flavonoids, tannins, catechins, vitamins C and E, and β-carotene may give better protection/bioactivity than single phytochemicals (Bazzano et al., 2003 2003 ). Phenolic compounds have been shown to have significant antioxidant effects (Newell et al., 2010 2010 ). Phenols, flavonoids, anthocyanins, and phenolic acids have been obtained from plants to attain an idyllic structural chemistry for scavenging free radicals. The antioxidant potential of polyphenols may arise from their great reactivity as hydrogen or electron donors. The polyphenol-derived radical stabilize and delocalize the unpaired electron (Chanda and Dave, 2009 2009 ). The scavenging of DPPH and ABTS radicals may be associated with the hydroxyl groups in caffeic acid and chlorogenic acid. The radical scavenging activity of phenolic compounds is influenced by the number of hydroxyl groups on the aromatic ring (Lazarova et al., 2014 2014 ). Both rutin and quercetin significantly chelate metal ions and have ferric reducing antioxidant power (Azuma et al., 1999 1999 ). According to Kim et al. ( 2004 2004 ), rutin and quercetin express their anti-inflammatory activity through modulation of the expression of pro-inflammatory genes such as cyclooxygenase-2 and inducible nitric oxide synthase, and via some vital cytokines. Based on these distinctive mechanisms, flavonoids are considered as plausible substances for anti-inflammatory activity. The presence of caffeic acid, chlorogenic acid, rutin, and quercetin has been reported in many pteridophytes, namely P . multifida , P . vittata , P . aquilinum , P . leucotomos , Cyathea phalerata , S . tamariscina , O . sensibilis , M . struthiopteris , D . linearis , P . nudum , E . arvense , and A . capillus-veneris (Table 1 1 ). Free radicals are generated in cells where they accumulate and react with macromolecules, like lipids, proteins, and nucleic acids, thereby disrupting cellular functions. In addition, oxidative damage to cellular components results in alteration of membrane properties such as fluidity, ion transport, enzyme activity, and protein cross-linking, causing cell death (Chauhan and Chauhan, 2006 2006 ). Bio-compounds containing antioxidant potential have the capacity to inhibit carcinogenesis (Wang et al., 2011 2011 ; Firdaus et al., 2013 2013 ). In earlier studies, apigenin, quercetin, and luteolin were shown to inhibit cancer cell proliferation (Mishra et al., 2013a 2013a , 2013b 2013b ; Fotsis et al., 1997 1997 ). Moreover, phenolics have been observed to postpone or inhibit the progression of various diseases by preventing peroxidation of lipid membranes (Kumar et al., 2013a 2013a , 2013b 2013b ). Some bioactivity, such as lipoprotective, antiplatelet, and anti-inflammatory activity, is closely associated with antioxidative flavonoids (Kumar et al., 2012 2012 ). Flavonoids have the potential to be reducing agents and to scavenge free radicals, chelate metal ions and inhibit enzymatic systems (topoisomerases and kinases) responsible for free radical generation (Kumar and Pandey, 2013 2013 ). Yang et al. ( 2013 2013 ) reported the inhibition of cancer cell lines, such as stomach cancer, prostate cancer, malignant melanoma, and mouse fibroblasts, by crude extracts and several isolated compounds (Table 1 1 ) of C . fortunei . In addition, apigenin was found to induce apoptosis and suppress insulin-like growth factor I receptor signalling in human prostate cancer (Way et al., 2004 2004 ; Shukla and Gupta, 2009 2009 ). Some flavonoids, such as quercetin, apigenin, kaempferol, and rutin in Murraya koenigii , inhibit endogenous 26S proteasome activity in MDA-MB-231 cells (Noolu et al., 2016 2016 ). Several authors have reported the presence of apigenin, kaempferol, and luteolin in following species, namely P . multifida , P . aquilinum , Selaginella sp., A . capillus-veneris , C . phalerata , P . vittata , O . sensibilis , M . struthiopteris , Equisetum ramoissimum , S . doederleinii , D . linearis , P. nudum , Pityrogramma tartarea , P. calomelanos , Cheilanthes concolor , Cardamine flexuosa , Cardamine goyazensis , Pellaea cymbiformis , P. gleichenioides , P. pinnata , P. riedelii , Pteris altissima , Pteris angustata , Pteris decurrens , Pteris deflexa , Pteris denticulata , Pteris plumula , Pteris podophylla , Pteris splendens , Pteris propinqua , and C . fortunei (Table 1 1 ). Synergism between several flavones and flavonols for antimicrobial activity was confirmed in an earlier study. Moreover, quercetin and naringenin were found to significantly inhibit bactericidal motility (Yilmaz and Toledo, 2004 2004 ). The release of ROS from activated neutrophils and macrophages causes inflammation injury. Excessive production may lead to tissue injury by damaging macromolecules and lipid peroxidation of membranes (Winrow et al., 1993 1993 ; Gutteridge, 1995 1995 ). ROS inflammation stimulates the release of cytokines, such as interleukin-1, tumour necrosis factor-α, and interferon-α, which stimulate the production of extra neutrophils and macrophages. However, free radicals are the main mediators promoting the inflammatory process. By neutralizing free radicals, antioxidants can reduce inflammation (Delaporte et al., 2002 2002 ; Geronikaki and Gavalas, 2006 2006 ). Therefore, we urge biologists to focus on the mechanisms of action of bioactive compounds from early tracheophytes (pteridophytes) in medicinal applications. Conclusions 1 Several researchers have studied the phytochemical and medicinal uses of lycophytes and ferns in recent years. These studies have demonstrated the importance of ferns and their allies in plant science fields. We have listed here the medicinal uses of pteridophytes, specifically their antioxidant, antidiabetic, anti-cancer, antiviral, anti-inflammatory, wound healing, antimicrobial, and anti-Alzheimer activity. No significant review papers on medicinal pteridophytes have been published recently, only a few articles with inadequate reports. The medicinal pteridophytes listed in this review belong to the following thirty families: Davalliaceae, Equisetaceae, Lygodiaceae, Ophioglossaceae, Polypodiaceae, Psilotaceae, Pteridaceae, Salviniaceae, Selaginellaceae, Tectariaceae, Woodsiaceae, Aspleniaceae, Athyriaceae, Blechnaceae, Cyatheaceae, Dennstaedtiaceae, Dicksoniaceae, Dryopteridaceae, Hymenophyllaceae, Lindsaeaceae, Thelypteridaceae, Marsileaceae, Nephrolepidaceae, Onocleaceae, Marattiaceae, Oleandraceae, Osmundaceae, Cystopteridaceae, Gleicheniaceae, and Huperziaceae. Our review might encourage an appreciation of the uses of pteridophytes beyond ornamental purposes and, in particular, promote their use and development for medicinal benefits. Future directions 1 Pteridophytes comprise around 13 000 species that belong to nearly 48 plant families, but only a small number of species from thirty families have been analyzed for their biological activity. Hence, we urge biologists to evaluate the medicinal uses of pteridophytes and their bioactive principles. Using biotechnological tools, the efficacy of bioactive compounds could be increased by studies of in vitro culture, biosynthetic pathways, modes of action, and genetic expression. In addition to studies of their biological activity, pharmacokinetics, and toxicological profiles, multidrug therapies should be developed from early tracheophytes and pteridophytes. Furthermore, the genetic relationships among pteridophytes should be analyzed based on their secondary metabolite synthesis. 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This article does not contain any studies with human or animal subjects performed by any of the authors. 10.1631/jzus.B1600344 Project supported by the Science and Technology Project for Fundamental Research of Shenzhen (No. JCYJ20140905095624296) and the Basic Work Special Project of the Ministry of Science and Technology of China (No. 2013FY111500) 2016-07-28 2017-03-05 2022-01-05 Copyright © Zhejiang University and Springer-Verlag GmbH Germany, part of Springer Nature 2018 2018 Journal of Zhejiang University-SCIENCE B (Biomedicine and Biotechnology) 02 19