Unique Bioactive Secondary Metabolites of Ferns and Their Enhancement by Abiotic Stress: Medicinal Potential and...

Ferns represent an evolutionarily distinct group of vascular plants and constitute an underexplored source of structurally diverse secondary metabolites with potential medicinal value. Several fern-derived compounds, including sesquiterpenes, triterpenes, flavonoids, phloroglucinol derivatives, lactones, and glycosides, have been associated with antibacterial, antidiabetic, analgesic, anticancer, hepatoprotective, neuroprotective, and other biological activities. However, despite their biochemical uniqueness and long-standing use in traditional medicine, ferns remain less extensively investigated than angiosperms as sources of bioactive compounds. In addition to their natural phytochemical diversity, the production of secondary metabolites in ferns may be influenced by abiotic stressors, such as light quality and intensity, temperature, salinity, drought, water availability, and mineral nutrition. Available studies indicate that selected abiotic stress conditions can enhance the accumulation of phenolic acids, flavonoids, polyphenols, carotenoids, and related compounds in several fern families, including Aspleniaceae, Athyriaceae, Dryopteridaceae, Onocleaceae, and Thelypteridaceae. Nevertheless, information on stress-induced modulation of metabolites that are unique or highly characteristic of ferns, particularly terpenes, terpene glycosides, and specific flavonoid derivatives, remains limited. This review summarizes the current knowledge on unique secondary metabolites in ferns, their reported medicinal properties, and the potential use of abiotic stress as an elicitation strategy to enhance their production. Overall, the review highlights ferns as promising but still insufficiently explored reservoirs of bioactive metabolites and identifies key directions for future phytochemical, pharmacological, and cultivation-based research.