Antifungal potential of marine macroalgae: an integrative reviewAbstract
The bioprospecting of compounds from marine macroalgae represents a significant trend in the development of antifungal agents to combat the impacts caused by pathogenic and mycotoxin-producing fungi. Accordingly, this study seeks to organize this evidence for the period between 2015 and 2025, utilizing several databases: CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior), Google Scholar, PubMed, ScienceDirect, and Web of Science. The selected articles were subsequently organized using Excel. Forty-six articles were included in the review, with the studies primarily focusing on the African (55.6%) and Asian (24.4%) continents. A total of 66 genera and 127 species of macroalgae were identified, with the genera Ulva (40%) and Sargassum (31%) appearing most frequently. Fifty-seven fungal species were also identified among those tested, with the most frequent genera being Candida (44.4%), Aspergillus (37.8%), and Fusarium (33.3%). Furthermore, the presence of terpenoid, fatty acids, tannins, steroids, polysaccharides, flavonoids, and other compounds were observed in the extracts obtained, while the techniques for isolating microorganisms and extracting their chemical composition were discussed. In summary, the findings highlight the biotechnological potential of marine macroalgae extracts as promising sources of antifungal compounds for pharmaceutical and agricultural applications.
Keywords:
antimicrobial potential; biotechnology; fungi; seaweed
Resumo
A bioprospecção de compostos de macroalgas marinhas configura uma tendência relevante para o desenvolvimento de agentes antifúngicos frente aos impactos causados por fungos patogênicos e produtores de micotoxinas. Diante disso, o presente estudo busca organizar essas evidências no período entre 2015 e 2025, onde foram utilizadas distintas bases de dados: Coordenação de Aperfeiçoamento de Pessoal de Nível Superior – CAPES, Google Scholar, PubMed, ScienceDirect e Web of Science. Seguidamente realizou-se a organização dos artigos selecionados no programa Excel. Foram incluídos 46 artigos na revisão, em que se observou como principal foco desses estudos os continentes africano (55,6%) e asiático (24,4%). Verificou-se 66 gêneros e 127 espécies de macroalgas utilizadas, onde destacam-se como mais recorrentes os gêneros Ulva (40%) e Sargassum (31%). Também foram identificadas 57 espécies de fungos testados, sendo os gêneros mais frequentes Candida (44,4%), Aspergillus (37,8%) e Fusarium (33,3%). Além disso, observou-se a presença de ácidos graxos, terpenoides, taninos, esteroides, polissacarídeos, flavonoides e demais compostos nos extratos obtidos, enquanto discutiu-se sobre as técnicas de isolamento dos microrganismos e extração da composição química. Em síntese, enfatiza-se o potencial biotecnológico dos extratos de macroalgas marinhas como fontes promissoras de compostos antifúngicos para aplicações farmacêuticas e agrícolas.
Palavras-chave:
potencial antimicrobiano; biotecnologia; fungos; algas marinhas
1. Introduction
The natural marine potential has received increasing attention over the years due to the wide range of bioactive compounds already reported in different organisms (Carroll et al., 2021; Hentati et al., 2020), such as microorganisms (Zha et al., 2024), mollusks (Xiang et al., 2022), algae (Nurkolis et al., 2023), as well as shells, aquatic worms, soft corals, and fish, among others (Lu et al., 2021). Therefore, studies that involve marine biotechnology through bioprospecting represent a relevant trend in innovation (Calle, 2017).
Regarding the activities derived from the rich marine environment, antiviral, antibacterial, and antifungal perspectives are particularly relevant (Carroll et al., 2020). The secondary metabolites responsible for these bioactive activities originate from defense mechanisms developed in response to ecological pressure such as territorial competition, predation impact, and tidal fluctuations. In this context, the antimicrobial activity of some compounds inhibits or limits the development of competing microorganisms (Pérez et al., 2016).
Marine macroalgae, belonging to the groups Phaeophyta, Rhodophyta, and Chlorophyta, stand out for their versatility and multiple applications, especially in agriculture, pharmaceuticals, cosmeceuticals, nutraceuticals, among others. This wide applicability largely results from the bioactive compounds produced by these organisms, such as polysaccharides, peptides, phlorotannins, phenols, carbohydrates, proteins, fatty acids, and smaller compounds such as sterols. Among the biological activities associated with these compounds, their antimicrobial potential stands out (Cabral et al., 2021; Leandro et al., 2020).
Current literature indicates the high potential of marine products as therapeutic agents for fungal infection treatment, and secondary metabolites isolated from algae rank among the most investigated in this context (El-Hossary et al., 2017). However, studies differ substantially in terms of collection sites, macroalgal species, extraction solvents and methodological approaches, including crude extracts, and in vitro or in vivo assays, directly influencing the reported bioactivities (Mohy El-Din and Mohyeldin, 2018; Rengasamy et al., 2020).
Fungi represent a significant proportion of eukaryotic species, interact with nearly all forms of life on the planet, and play major roles in global ecosystems (Niskanen et al., 2023). However, this broad presence increases the potential emergence of fungal infections that may affect agriculture, the environment, and animal and human health, generating significant social and economic impacts. Candidiasis and aspergillosis exemplify diseases that pose serious threats to public health, alongside other emerging conditions that have become contemporary concerns. (Almeida et al., 2019; Lass-Flörl and Steixner, 2023).
Fungal infection, or contamination caused by pathogenic fungi and/or mycotoxin producers, has driven the search for effective fungicides to control these agents. However, the continuous use of chemical compounds is associated with resistant strains and negative environmental impacts. Therefore, biopreservation emerges as a promising strategy by integrating the search for sustainable alternatives that ensure food safety and contribute to new drug development. Within this scenario, macroalgae potential appears as an innovative approach to manage these infections by offering bioactive compounds with relevant antifungal properties (Agarwal et al., 2021; Sánchez-Torres, 2025). This review aims to discuss the antifungal properties of macroalgae worldwide, with emphasis on applied methodologies and the main results reported in the literature.
2. Method
The bibliographic survey was conducted in February 2025 using the CAPES Journals Portal (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES, Brazil) as the primary data source, in addition to Google Scholar, PubMed, ScienceDirect, and Web of Science. The following descriptors were applied: “macroalgae” OR “seaweed” OR “algae” AND “antifungal activity” OR “antimicrobial” AND “active compounds” AND “extracts”.
The selected studies covered the period from 2015 to February 2025. Inclusion criteria comprised articles published in English that reported methodologies involving macroalgae collection, extract preparation, and in vitro application against fungal strains. Exclusion criteria included review articles or book chapters, studies evaluating extracts against microorganisms other than fungi, and investigations based exclusively on in vivo assays; this was due to the limited number of available in vivo investigations and their marked methodological heterogeneity, allowing for greater comparability among the selected studies.
Subsequently, the selected studies were compiled and organized using Microsoft Excel, allowing categorization according to macroalgal and fungal genera and species, types of extract preparation, microbial testing methodologies, observed effects, identified compounds, and publication information (authors, journals, and year of publication). These data were organized into tables and figures to facilitate comparative analysis, characterizing the present study as an integrative review (Souza et al., 2010).
3. Results
After the article search, we identified 8,960 records across the databases: CAPES (177), Google Scholar (8,400), PubMed (42), Science Direct (337), and Web of Science (4). After the initial screening by titles and abstracts, according to the established inclusion and exclusion criteria, we read 251 articles and selected 70. We then removed 24 duplicates, which resulted in a total of 46 articles included in the review.
Over the ten-year period analyzed, publications increased markedly between 2021 and 2024, which account for approximately 46.6% of the total articles included in the analysis (Figure 1A). Furthermore, according to (Figure 1B), Africa recorded the highest number of studies in the field (55.6%), followed by Asia (24.4%), Europe (8.9%), North America (6.7%), and South America (4.4%). At the country level, Egypt stands out with the largest publication volume (26.66%), followed by India (13.33%).
Temporal and geographical distribution of selected articles. (A) Percentage of articles identified by year of publication; (B) Percentage of studies by continent.
We identified 127 macroalgal species distributed among three phyla: Rhodophyta (41.73%), Phaeophyta (32.28%), and Chlorophyta (25.98%). As shown in Figure 2, we recorded 66 genera in total, with Ulva sp. (40%) among green algae and Sargassum sp. (31.1%) among brown algae as the most recurrent. The species Ulva lactuca and Sargassum vulgare were the most representative within their respective groups in both diversity and frequency. Among red algae, the genera Gelidium and Hypnea showed the highest frequency, each accounting for 13.3% of the records studies analyzed.
Percentage of articles classified by genus of macroalgae tested from an antifungal perspective.
Regarding the fungal species used in tests with macroalgal extracts, we identified 57 species belonging to 27 distinct genera (Figure 3A).
Most frequent fungal genera and species reported in the studies. (A) Percentage of articles by fungal genus; (B) Percentage by species within the genus Candida; (C) Percentage by species within the genus Aspergillus.
Considering the recurrence of fungal genera in the analyzed studies, Candida (44.4%), Aspergillus (37.8%), Fusarium (33.3%), and Penicillium (28.9%) were the most frequently investigated. Among the most recurrent species, Candida albicans appeared in 17 studies (37.8%) and Aspergillus niger in 12 studies (26.7%), as shown in Figures 3B and 3C. The Genera Alternaria sp. (15.5%), Botrytis sp. (15.5%), and Mucor sp. (6.6%) also stood out, especially the species Alternaria alternata and Botrytis cinerea, as well as Mucor spp. (Figure 3).
Regarding the main biotechnological or industrial applications explored in studies with marine macroalgae, research focused predominantly on the medical, pharmaceutical, agricultural, and food sectors. Among these, studies aimed at improving crop agricultural quality were the most frequent. In addition, several investigations addressed more than one application sector, emphasizing fungi associated with biofilm formation or responsible for respiratory diseases, skin infections, diabetes, and other disorders, as well as phytopathogens affecting roots, leaves, and fruits.
Concerning extract production, studies employed different preparation methodologies, solvents, and dosages. The most common extraction techniques included solvent maceration, Soxhlet extraction, sonication, centrifugation, and percolation. Regarding solvents used in maceration or partition processes, the most frequent were water, ethanol, methanol, ethyl acetate, hexane, acetone, dichloromethane, chloroform, among others (Table 1), at concentrations ranging from 15.6 µL/mL to 1000 µL/mL.
Overview of extraction methods, antifungal assays, and main compounds identified in marine macroalgae extracts.
In general, tests conducted with macroalgae demonstrated significant antifungal activity, ranging from moderate effects (inhibition zones ranging from 12 to 60 mm) to complete inhibition of fungal growth. Only a small proportion of studies (4.4%) did not report satisfactory responses. Analyses mainly evaluated morphological alterations and the action of these products on parameters such as fungistatic effect, inhibition zone formation, minimum inhibitory concentration (MIC), minimum fungicidal concentration (MFC), and inhibition of spore germination. For this purpose, researchers applied different methodologies, including disk diffusion assays, agar well diffusion, poisoned food technique, and broth dilution or microdilution tests, as described in Table 1.
Among the analyzed studies, 73.3% also investigated the chemical composition of the extracts using analytical methodologies such as GC-MS, HPLC, LC-HR-ESI-MS, UV–Vis spectroscopy, and phytochemical screening. Additional methods, including FTIR-ATR, DPPH assays, SDS-PAGE and circular dichroism, were also employed to identify and characterize bioactive compounds.
The results predominantly revealed the presence of different classes of bioactive compounds, such as fatty acids (30.3%), terpenoids (24.2%), tannins (21.2%), steroids (15.1%), polysaccharides (12.1%), flavonoids (9.1%), polyphenols/phenols (9.1%), proteins (9.1%), hydrocarbons (9.1%), alkaloids (6.1%), terpenes (6.1%), carbohydrates (3.0%), among others (Figure 4).
4. Discussion
Marine macroalgae have gained increasing attention as sources of bioactive molecules. Ghaliaoui et al. (2024) detailed biocompounds extracted from macroalgae containing minerals, amino acids, proteins, fatty acids, lipids, polysaccharides, dietary fibers, vitamins, and several secondary metabolites such as phenols, alkaloids, terpenes, and pigments, many of which exhibit antifungal properties.
Consistently, the temporal analysis of the selected studies indicates a significant increase in scientific publications between 2021 and 2024, which concentrated 46.6% of the analyzed articles. This increase reflects growing scientific interest in marine biotechnology and natural metabolites. The predominance of African (55,6%) and Asian (24,4%) continents in macroalgal bioprospecting research can be explained by several factors.
Pradhan et al. (2020) state that seaweeds are widely used in diet and traditional medicine in Asian countries due to the presence of essential nutrients. Asian and African regions present high potential for exploration and cultivation of marine algae, including scientifically underexplored species, due to the extensive biodiversity of their coastal zones (Akbar and Hasan, 2024; Msuya et al., 2022). In addition, investment in blue biotechnology research has intensified in developing countries, especially in coastal regions economically dependent on marine resources (Bax et al., 2022; Selig et al., 2019). In Egypt, for example, 12 articles on these bioprospecting studies were identified, representing 26% of the total. This finding aligns with the country’s strategic location between the Mediterranean and Red Seas and with the activity of Egyptian institutions in prospecting antifungal compounds from macroalgae (El-Sheekh et al., 2022). Other prominent countries include India (6), Morocco (3), and Mexico (3).
Conversely, in the South American context, despite high biodiversity, the region still does not stand out among major hubs of marine-resource-based biotechnological applications. This scenario results from limited scientific exploration, insufficient infrastructure, and the absence of consolidated protocols for studying these metabolites. In Brazil, megadiversity offers opportunities for biotechnological development. Regarding marine algae utilization, stronger integration among different knowledge areas with a multidisciplinary approach becomes essential to advance the generation and application of this information (Santos et al., 2023; Quintana-Manotas et al., 2024).
Among the macroalgae most frequently identified in the studies, the genera Sargassum and Ulva stand out. Besides being among the most extensively studied macroalgae, their antifungal activity has been associated with the combination of bioactive compounds and the environmental conditions to which they are subjected (Pérez et al., 2016).
Regarding the microorganisms studied, Candida sp., Penicillium sp., and Aspergillus sp. corresponded to 35.08% of the studies analyzed. Penicillium and Aspergillus represent major mycotoxigenic fungi in the human food chain, whereas Candida species rank among the most common human fungal pathogens responsible for superficial and systemic infections (Sweeney and Dobson, 1998; Turner and Butler, 2014).
The increased use of antifungal agents in both medical therapy and agriculture has contributed to resistance against drugs such as triazoles and echinocandins, especially in Aspergillus and Candida species. Resistance may arise from reduced intracellular drug concentration, genetic alterations in enzymatic targets, changes in membrane composition, and inefficient compound activation (Gonçalves et al., 2016; Vanden Bossche, 1997).
Additionally, studies face many complexities in the standardization and scientific validation of biological controls for pathogens. Choosing extraction and isolation methods for natural products is essential due to the diversity of available techniques and purposes. Researchers must carefully evaluate objectives, temperature, and extraction time (Quitério et al., 2022). Among the analyzed studies that used positive and negative controls, maceration remained the most used extraction method, followed by Soxhlet extraction, and in some cases multiple techniques were combined to enhance extraction efficiency. Furthermore, methanol, ethanol, and water were the most common solvents, although several studies used more than one solvent.
Another fundamental step in investigating marine algal biocompounds is defining purification, identification, and validation techniques (Cermeño et al., 2020). As observed in the studies analyzed, secondary metabolite groups such as fatty acids, terpenoids, phenolics, flavonoids, and steroids predominated, and several macroalgal studies indicate these classes as promising antifungal agents. Guimarães and Venâncio (2022) highlight the significant antifungal and mycotoxin-inhibitory potential of fatty acids, mainly through membrane destabilization, enzyme inhibition, and metabolic pathway interference. Konuk and Ergüden (2020) emphasize the ability of terpenoids to damage plasma membranes and inhibit fungal growth by penetrating the lipid bilayer, resulting in ion leakage and altered membrane pH and conductivity. The review by Davidova et al. (2024) describes the broad antifungal spectrum of polyphenols, which disrupt membrane integrity and permeability and induce oxidative effects that compromise essential microbial metabolism.
Marine macroalgae contains a wide diversity of bioactive compounds with antifungal activity, although most studies remain limited to in vitro assays. Beyond the fact that research efforts have focused on specific genera, this activity is directly linked to the combination of compounds present in the macroalgae and the environmental conditions under which they are applied. Despite scientific advances, the limited number of in vivo studies, the lack of methodological standardization, and data gaps regarding regions of high biodiversity, such as South America, continue to limit the consolidation of available evidence.
5. Conclusion
The studies analyzed indicate that marine macroalgae have been increasingly investigated for their antifungal properties. African and Asian regions stand out in this scenario, whereas South America shows the most significant research gaps. Studies focus mainly on the medical, pharmaceutical, agricultural, and food sectors, particularly on phytopathogen control. The diversity of solvents and methodologies, together with the identification of chemical groups such as fatty acids, terpenoids, tannins, steroids, polysaccharides, flavonoids, and polyphenols, reinforces the broad range of algal properties. Despite limitations noted in the literature regarding the predominance of in vitro over in vivo studies, as well as methodological heterogeneity, the results of this review highlight growing scientific interest in the potential of marine macroalgae as sources of antifungal compounds. Therefore, this study provides an important basis for the development of future research in this field.
Data Availability Statement
The research data analyzed in this study are not publicly available in any database.
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