Abstract
Usually regarded as less evolved than their more recently diverged vascular sisters, which currently dominate vegetation landscape, bryophytes seem having nothing to envy to the defensive arsenal of other plants, since they had acquired a suite of chemical traits that allowed them to adapt and persist on land. In fact, these closest modern relatives of the ancestors to the earliest terrestrial plants proved to be marvelous chemists, as they traditionally were a popular remedy among tribal people all over the world, that exploit their pharmacological properties to cure the most different diseases. The phytochemistry of bryophytes exhibits a stunning assortment of biologically active compounds such as lipids, proteins, steroids, organic acids, alcohols, aliphatic and aromatic compounds, polyphenols, terpenoids, acetogenins and phenylquinones, thus it is not surprising that substances obtained from various species belonging to such ancestral plants are widely employed as antitumor, antipyretic, insecticidal and antimicrobial. This review explores in particular the antifungal potential of the three Bryophyta divisions—mosses (Musci), hornworts (Anthocerotae) and liverworts (Hepaticae)—to be used as a sources of interesting bioactive constituents for both pharmaceutical and agricultural areas, providing an updated overview of the latest relevant insights.
Highlights
Plants stand as an infinite resource for drug development: it has been estimated that the 11% of the 252 drugs considered as basic and essential by the WHO are obtained from flowering plants [2] but a wide amount of phytochemicals provided with antimicrobial properties was characterized and isolated in representatives of each taxa, from algae to pteridophytes to higher plants; bryophytes, non-vascular early land plants considered the link between seed and vascular plants to their algal ancestors, contribute to this “phytochemical treasure chest”
Among secondary metabolites involved in abiotic stress, terpenoids have long time been recognized to play an important role in bryophyte, and in vascular plants, environmental interactions [5,136]: over the last four decades, more than 1600 compounds belonging to the terpenoids family have been reported from this plant lineage [6,42,60], a number of them demonstrating the ability to possess antioxidant, antimicrobial and cytotoxic activities that suggest a potential for pharmaceutical purposes—mainly addressed to emerging drug resistance occurrences
Humankind has always been interested in plants as a source of food, and as a tool for healing purposes; since the herbal medicine represented the first and foremost therapeutic tool available to humans for many centuries, the history of plant-derived drugs—starting from Dioscoride’s De materia medica—is as old as the world
Summary
As masterfully reviewed by Davey and Currah [32], various types of association, both pathogenic and beneficial, occur in the “mosses chamber” of the bryophyte’s world In terms of their development, these interactions clearly differ from those occurring between fungi and higher plants, since characteristic targets for predation (such as storage organs and specialized nutrient transport tissues) and entrance (regulated stomata and lignified, reinforced cell walls around vessels) are likely missing in bryophytes: the exploitation of degradative enzymes secreted from the penetration peg seems to represent the most common entry-ticket strategy, for example in Sphagnum pathogenic fungi [33,34,35]. It could be speculated that these compounds (namely polyphenols, flavonoids and anthocyanins), providing an ancestral layer of biochemical defenses in mitigating pathogen infection in liverworts, played a critical role for the terrestrialization of plants
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