• Open Access
    Mini Review

    Mini-review on the antimicrobial potential of actinobacteria associated with seagrasses

    Galana Siro 1*
    Atanas Pipite 2

    Explor Drug Sci. 2024;2:117–125 DOI: https://doi.org/10.37349/eds.2024.00038

    Received: August 10, 2023 Accepted: December 01, 2023 Published: February 29, 2024

    Academic Editor: Juergen Reichardt, James Cook University, Australia

    This article belongs to the special issue Exploring Potential Drugs from Natural Products


    The search for novel therapeutic agents to combat the crisis of antimicrobial resistance has spanned from terrestrial to unique, marine environments. Currently, most of the drugs available for usage are derived from microbial metabolites, especially those belonging to the bacterial group, actinobacteria. Actinobacteria are hotspot organisms that exist in all habitats with a myriad of unique biosynthetic metabolites. Seagrasses appear to be a key ecosystem within the coastal environment worth bioprospecting for novel natural products. Unfortunately, literature about the bioactive potential of their associated prokaryotes, including actinobacteria remains limited. In this context, this review focused on actinobacteria with antibiotic-producing capabilities derived from different parts of seagrass plants (i.e. roots, rhizomes, and leaves). To date, there were no purified molecules derived from seagrass-associated actinobacteria that were subjected to structure elucidation. From the underpinning of numerous biological profiles such as antibacterial, antifungal, and algicidal activities of seagrass-derived actinobacteria reported in this review during the period from 2012–2020, it provides a continual growth of knowledge accruing overtime, providing a foundation for future research.


    Actinobacteria, seagrasses, biosynthetic metabolites, marine ecosystem, antimicrobial resistance


    The crisis of multidrug resistance has created havoc in the health sector, with increasing ramifications for people’s health. The quantity of resistant genes differs geographically, thus requiring global attention to counteract this inevitable phenomenon [1, 2]. As a counter approach, the administration of antibiotics and the tedious bioprospects of novel bioactive compounds from marine sources are common [3, 4]. A quantum of information regarding the natural products from marine environments has received great recognition due to their diverse applications and efficiency [5]. Bioprospects in marine habitats have continuously provided a series of novel and interesting metabolites [68]. An important marine ecosystem represented in this review is seagrasses. Seagrass meadows support gradation in species diversity and high biodiversity of marine life [9]. Actinobacteria sometimes referred to as actinomycetes are Gram-positive bacteria and form a unique group of bacterial lineages. They belong to the phylum Actinomycetota (https://lpsn.dsmz.de/phylum/actinomycetota) and are regarded as hotspots for clinical drug discovery. Their genomes have a high guanine-cytosine (GC) content and genes associated with antibiotic production [1012]. The genus Streptomyces is the most widely gifted group of bacteria with efficient antibiotic machinery and is responsible for the high availability of clinical antibiotics [13, 14]. Actinomycetes exist as unicellular and multicellular, but some species function as either symbionts or pathogens. In spite of their antibiotic-producing capabilities, actinomycetes have other important functions including nutrient turnover and bioremediation processes [15, 16]. Moreover, actinomycetes are ubiquitous due to their adaptive nature. In tandem with this, they are able to colonize the oceans heterogeneous habitats, including seagrass meadows [1719]. Studies that focus on the isolation of bacteria producing antagonistic metabolites from seagrasses are limited. In the search for potential prospecting fields, this mini-review herein scrutinized actinobacteria associated with seagrasses for their pharmaceutical compounds. Data reported in this paper were obtained from Web of Science, Google Scholar, and other online databases using their respective advanced search option with the following search terms: “actinobacteria-derived from seagrass: metabolites and bioactivity”. In total, this review is an attempt to give more attention and momentum to seagrass-based actinobacteria and their medicinal active compounds.

    Seagrass biology, evolution, and biodiversity

    Seagrasses are submerged, marine flowering plants that thrive in the ocean except in the polar regions [20]. They comprise over 60 species, belonging to four exclusive plant families including Zosteraceae, Hydrocharitaceae, Posidoniaceae, and Cymodoceaceae, and support immense levels of biodiversity [2125]. Seagrasses have a range of features that arose from adapting to living in submerged marine conditions [26, 27]. As an aquatic angiosperm, their multiple colonization events in the ocean predated over 70 million years ago [25]. Notably, seagrasses are rhizomatous plants adapted to aquatic environments [28]. Depending on their growth sizes, seagrasses range from small species (e.g., Halophila engelmannii) to large species (e.g., Posidonia australis) [23]. The highly productive nature of seagrasses is largely influenced by a mixture of sexual and asexual clonal reproduction [26]. Collectively, these foundation species form dense meadows, flowering and seeding underwater with significant provisions that ripple through the entire coastal ecosystems. Seagrass meadows are part of the tropical seascape, similar to mangroves and coral reefs, but they have a wide distribution along both tropical and temperate coasts. All these important marine ecosystems are interconnected via biogeochemical processes. Among the triad of key marine habitats, seagrass meadows are regarded as one of the most productive and crucial components of the marine ecosystem. They act as nutrient cyclers, sediment stabilizers (coastal protection), organic carbon producers (carbon sequestration), and juvenile nursery grounds and feeding areas for marine life (provide trophic subsidy to contiguous habitats) [24, 28, 29]. The ecological importance of seagrass meadows is irrefutable, and yet remains poorly studied, in terms of medicinal properties [30]. Seagrass meadows straddle the subtidal and intertidal zones of the temperate and tropical coastlines. Their distribution is mostly restricted to regions where wave activity is limited, and there is sufficient light and nutrient availability [31]. The capacity and performance of seagrasses are directly proportional to their area of coverage and density. Seagrass meadows show remarkably high levels of primary productivity and exist as monospecific or multispecies communities, depending on the bioregions [28]. Its mass coverage spans from small patches to extensive underwater lawns [28].

    The ocean often receives a cocktail of raw sewage or pollutants. Consequently, shoreline microbial populations can spike to dangerous levels with serious health implications [32]. As one of the many natural counter approaches, seagrass plants often have innate filtration and responsive systems that facilitate the attenuation of nutrient levels, microplastic levels, microbial populations, and turbidity [3336]. In most cases, it involves the deposition process, where they ensnare particulates and microbes drifting through the ocean by ensuring affable conditions to a certain degree. Despite their critical values, the fragmentation of seagrass meadows involves both natural pressures (waves, currents, and extreme weather events) and anthropogenic pressures (dredging and infilling, recreational activities, raw sewage discharges, eutrophication, and coastal constructions) [37]. A significant feature of seagrass meadows is the various constituency of the seagrass holobiont. Seagrasses, similar to terrestrial plants, house communities of microbes, including actinobacteria that exhibit symbiotic relations. Moreover, the seagrass microbiome consists of intricate interactive networks that facilitates their overall fitness and growth [26, 3841]. Both the seagrass host and their associate microbes are capable of producing potent metabolites that prevent the invasion of opportunistic pathogens [39, 42]. Studies have shown that distinct microbial communities exist in discrete microenvironments of seagrasses [43]. Overall, the seagrass microbiome differs in composition between the different plant parts (i.e. roots, rhizomes, and leaves), as well as between species, which have a vast geographical distribution and are subject to a variety of environmental conditions. There are also differences between seagrass microbiomes and those of the adjacent seawater and surrounding sediment [4447].

    Traditional and medicinal use of seagrass around the world

    There is evidence throughout history of the use of seagrass as food, medicine, fertilizer, and livestock feed [20, 48, 49]. In many places, such as the village of Chwaka in East Africa, they have a mixed economy that relies heavily on seagrasses beds which provide cash income and the most important source of daily protein (fish associated with seagrasses) [20]. India is another region of the world where seagrasses form an important part of the local economy. A study has shown that the information on the nutritional value of seagrass has been found to be equivalent to that of Bengal gram, peas, potatoes, and southern potatoes and is completely safe for consumption by analyzing the concentrations of toxic elements: lead (Pb), chromium (Cr), and cadmium (Cd) [20]. In Tunisia, the leaves of the seagrass, Posidonia oceanica, which have antifungal and insect repellent properties, have been used as livestock bedding and as feed supplements for poultry and livestock [48]. In most European and Mediterranean coastal countries, seagrasses have been used for different purposes such as packaging equipment for transporting fragile items (i.e. glassware, pottery) to ship fresh fish from the coast to cities, bedding for livestock in stables, filler for mattresses and cushions (respiratory infections seemed to be prevented from sleeping in this type of bedding), roof insulation (i.e. in southeastern Spain and the Balearic Islands), and also a roof cover (i.e. in Netherlands) [49].

    The use of seagrass throughout history has shown that there is knowledge of the wide range of qualities possessed by these oceanic plants [50]. However, with the improvement of science and technology, studies have shown that these properties are based on the fact that seagrasses themselves can produce valuable chemical compounds or that these chemical compounds can be produced by microorganisms in symbiotic relationships with these seagrasses [50]. Remarkably, a total of 154 natural compounds derived from 70 seagrass species have been reported so far, predominantly from the host seagrass itself [50].

    Diversity and bioactive profiles of seagrass-derived actinobacteria

    The isolation of marine actinobacteria is influenced by the isolation parameters such as culture, pondus hydrogenii (pH) and temperature, incubation time, and concentration of the medium [51]. Numerous studies attest to culture seagrass-derived actinobacteria by employing the cultivation parameters as follows: using a wide variety of media, the addition of natural seawater, artificial seawater, or deionized/distilled water with different concentrations of sodium chloride, culture temperature 26°–29°C, and an incubation time of 1–6 weeks [5257]. Certain actinobacterial have been isolated from all seagrass parts (roots, rhizomes, and leaves), whereas, most research specifically targeted actinobacteria as endophytes from the roots and leaves of seagrass, as shown in Table 1.

    Genus/species of actinobacteria isolated from seagrasses between 2012 and 2020

    Genus/speciesFamilySeagrass species and nature of sampleBioactivityCountryReference
    Streptomyces spp., Micromonospora spp., Verrucosispora (Micromonospora) sp., Saccharomonospora spp., Actinomycetospora sp., Microbacterium sp., Mycobacterium spp., Nonomuraea sp., Nocardiopsis sp., and Glycomyces sp.Streptomycetaceae, Micromonosporaceae, Pseudonocardiaceae, Microbacteriaceae, Mycobacteriaceae, Streptosporangiaceae, Nocardiopsidaceae, and GlycomycetaceaeThalassia hemprichii (whole plant)Contain nonribosomal peptide synthetase (NRPS) and polyketide synthase (PKS) genes (antibacterial activity)China[52]
    Streptomyces spp.StreptomycetaceaeSyringodium isoetifolium (leaves and roots)Antibacterial activityIndia[53]
    Saccharomonospora sp., Kocuria sp.Pseudonocardiaceae, MicrococcaceaeCymodocea serrulate (roots)Phosphate solubilizing, nitrogen-fixing ability, and enzyme activityKasuwari Island, India[54]
    Arthrobacter spp.MicrococcaceaeZostera marina and Zostera japonica (leaves)Algicidal activityPuget Sound, USA[73]
    Kocuria palustris, Kocuria atrinae, Arthrobacter flavus, Ornithinimicrobium humiphilum, Corynebacterium afermentans subsp. afermentansMicrococcaceae, Ornithinimicrobiaceae, and CorynebacteriaceaeHalodule uninervis (soil and root)Antifungal activitySaudi Arabia[55]
    Not availableCellulomonadaceae, MicrobacteriaceaeHalophila ovalis (roots)No bioactive test performedAustralia[74]
    Streptomyces sp.StreptomycetaceaeCymodocea rotundata (whole plant)Antimicrobial activityIndonesia[71]
    Streptomyces lienomyciniStreptomycetaceaeEnhalus acoroides (leaves)Antibacterial activityIndonesia[57]
    Isoptericola sp., Rhodococcus sp., and Streptomyces spp.Promicromonosporaceae, Nocardiaceae, and StreptomycetaceaeZostera marina (leaf and associate sediment)No bioactive test performedBodega Bay, USA[56]
    Display full size

    Seagrasses harbor a rich pool of specialized metabolites [58, 59]. The antimicrobial assay and compound elucidation of seagrass extracts have been examined under different contexts. In most cases, the results have shown that cytotoxic, antimicrobial, antimalarial, antioxidant, antibiofilm, anti-inflammatory, or antimicrofouling activities are prevalent and widespread among various solvent extracts from seagrass species [25, 6065]. In marine habitats, available surfaces are rapidly colonized by a spectrum of microbes. These microbial communities are often influenced by seasonality, and vegetative and non-vegetative sites [66]. With that, the vegetative cover of seagrasses provides a significant substratum for the rich diversity of organisms, including actinobacteria which forms an integral part of the seagrass ecosystem [9, 6769]. Actinobacteria are among the pioneer colonizers of seagrasses [70] and their diversity varies between seagrass species [52, 53]. The seagrass-derived actinomycetes are capable of producing antimicrobial agents (Table 1). At the time of this review, there were no purified molecules derived from seagrass-associated actinobacteria that were subjected to structure elucidation. Studies have shown that they have a spectrum of biological potentials such as antibacterial activity [52, 53, 57, 71, 72], antifungal activity [55, 71], algicidal activity [73], nitrogen-fixing ability, and enzymatic activity [54]. Conspicuously, there are available studies where no bioactive test was performed for the actinobacteria discovered [56, 74]. Interestingly, Wu et al. [52] reported the isolation of Verrucosispora sp. from the seagrass Thalassia hemprichii (this was the first time a Verrucosispora strain was isolated as a plant endophyte). The members of the genus Verrucosispora have subsequently all been transferred to the genus Micromonospora (https://lpsn.dsmz.de/genus/verrucosispora).

    Concurrently, they also cultivated four potential novel strains of actinomycetes which are yet to be categorized. Overall, as studies multiply in this area, more detailed discoveries of promising metabolites will be unveiled.


    The seagrass meadows harbor a high biodiversity with a great wealth of specialized metabolites. The use of seagrass throughout history has shown that there is knowledge of the wide range of qualities possessed by these oceanic plants [50, 58]. This review highlights the tremendous bioactive potential of actinobacteria isolated from various species of seagrasses. Most available studies focus on seagrass extracts for antimicrobial assays, with limited attention based on their microbiome bioactivity. Future research should target seagrass meadows for actinobacteria-derived compounds. With the advent of improved genomic technologies, genome-based research focusing on the latter is important. Overall, the field of research targeting microbial biosynthetic potential associated with seagrasses is burgeoning.


    Author contributions

    GS: Conceptualization, Writing—original draft, Writing—review & editing. AP: Conceptualization, Writing—original draft, Writing—review & editing, Supervision. All authors read and approved the submitted version.

    Conflicts of interest

    The authors declare that they have no conflicts of interest.

    Ethical approval

    Not applicable.

    Consent to participate

    Not applicable.

    Consent to publication

    Not applicable.

    Availability of data and materials

    Not applicable.


    Not applicable.


    © The Author(s) 2024.


    Munk P, Brinch C, Møller FD, Petersen TN, Hendriksen RS, Seyfarth AM, et al.; Global Sewage Surveillance Consortium; Larsson DGJ, Koopmans M, Woolhouse M, Aarestrup FM. Genomic analysis of sewage from 101 countries reveals global landscape of antimicrobial resistance. Nat Commun. 2022;13:7251. Erratum in: Nat Commun. 2023;14:178. [DOI] [PubMed] [PMC]
    Brinch C, Leekitcharoenphon P, Duarte ASR, Svendsen CA, Jensen JD, Aarestrup FM. Long-term temporal stability of the resistome in sewage from Copenhagen. mSystems. 2020;5:e00841-20. [DOI] [PubMed] [PMC]
    Voser TM, Campbell MD, Carroll AR. How different are marine microbial natural products compared to their terrestrial counterparts? Nat Prod Rep. 2022;39:719. [DOI] [PubMed]
    Sigwart JD, Blasiak R, Jaspars M, Jouffray JB, Tasdemir D. Unlocking the potential of marine biodiscovery. Nat Prod Rep. 2021;38:123542. [DOI] [PubMed]
    Carroll AR, Copp BR, Davis RA, Keyzers RA, Prinsep MR. Marine natural products. Nat Prod Rep. 2022;39:112271. [DOI] [PubMed] [PMC]
    Siro G, Pipite A, Christi K, Srinivasan S, Subramani R. Marine actinomycetes associated with stony corals: a potential hotspot for specialized metabolites. Microorganisms. 2022;10:1349. [DOI] [PubMed] [PMC]
    Siro G, Donald L, Pipite A. The diversity of deep-sea actinobacteria and their natural products: an epitome of curiosity and drug discovery. Diversity. 2023;15:30. [DOI]
    Khan N, Yılmaz S, Aksoy S, Uzel A, Tosun Ç, Kirmizibayrak PB, et al. Polyethers isolated from the marine actinobacterium Streptomyces cacaoi inhibit autophagy and induce apoptosis in cancer cells. Chem Biol Interact. 2019;307:16778. [DOI] [PubMed]
    Borowitzka MA, Lavery PS, Keulen M. Epiphytes of seagrasses. In: Larkum AWD, Orth RJ, Duarte CM, editors. Seagrasses: biology, ecology and conservation. Dordrecht: Springer Netherlands; 2006. pp. 441–61.
    Mast Y, Stegmann E. Actinomycetes: the antibiotics producers. Antibiotics. 2019;8:105. [DOI]
    Barka EA, Vatsa P, Sanchez L, Gaveau-Vaillant N, Jacquard C, Meier-Kolthoff JP, et al. Taxonomy, physiology, and natural products of actinobacteria. Microbiol Mol Biol Rev. 2015;80:143. Erratum in: Microbiol Mol Biol Rev. 2016;80:iii. [DOI] [PubMed] [PMC]
    Anandan R, Dharumadurai D, Manogaran GP. An introduction to actinobacteria. In: Dhanasekaran D, Jiang Y, editors. Actinobacteria. Rijeka: IntechOpen; 2016. [DOI]
    Watve MG, Tickoo R, Jog MM, Bhole BD. How many antibiotics are produced by the genus Streptomyces? Arch Microbiol. 2001;176:38690. [DOI] [PubMed]
    Donald L, Pipite A, Subramani R, Owen J, Keyzers RA, Taufa T. Streptomyces: still the biggest producer of new natural secondary metabolites, a current perspective. Microbiol Res. 2022;13:41865. [DOI]
    Verma S, Kuila A. Bioremediation of heavy metals by microbial process. Environ Technol Innov. 2019;14:100369. [DOI]
    Sharma M, Dangi P, Choudhary M. Actinomycetes: source, identification, and their applications. Int J Curr Microbiol Appl Sci. 2014;3:80132.
    Singh R, Dubey AK. Diversity and applications of endophytic actinobacteria of plants in special and other ecological niches. Front Microbiol. 2018;9:1767. [DOI] [PubMed] [PMC]
    Jagannathan SV, Manemann EM, Rowe SE, Callender MC, Soto W. Marine actinomycetes, new sources of biotechnological products. Mar Drugs. 2021;19:365. [DOI] [PubMed] [PMC]
    Lam KS. Discovery of novel metabolites from marine actinomycetes. Curr Opin Microbiol. 2006;9:24551. [DOI] [PubMed]
    Rengasamy RRK, Radjassegarin A, Perumal A. Seagrasses as potential source of medicinal food ingredients: nutritional analysis and multivariate approach. Biomed Prev Nutr. 2013;3:37580. [DOI]
    Unsworth RKF, Nordlund LM, Cullen-Unsworth LC. Seagrass meadows support global fisheries production. Conserv Lett. 2019;12:e12566. [DOI]
    Duffy JE. Biodiversity and the functioning of seagrass ecosystems. Mar Ecol Prog Ser. 2006;311:23350. [DOI]
    Papenbrock J. Highlights in seagrasses’ phylogeny, physiology, and metabolism: What makes them special? Int Sch Res Not. 2012;2012:103892. [DOI]
    Orth RJ, Carruthers TJB, Dennison W, Duarte CM, Fourqurean JW, Heck KL, et al. A global crisis for seagrass ecosystems. Bioscience. 2006;56:98796. [DOI]
    Jeyapragash D, Saravanakumar A, Yosuva M. Seagrass metabolomics: a new insight towards marine based drug discovery. In: Zhan X, editor. Metabolomics. Rijeka: IntechOpen; 2021.
    Ugarelli K, Chakrabarti S, Laas P, Stingl U. The seagrass holobiont and its microbiome. Microorganisms. 2017;5:81. [DOI] [PubMed] [PMC]
    Sandoval-Gil JM, Ruiz JM, Marín-Guirao L. Advances in understanding multilevel responses of seagrasses to hypersalinity. Mar Environ Res. 2023;183:105809. [DOI] [PubMed]
    Short F, Carruthers T, Dennison W, Waycott M. Global seagrass distribution and diversity: a bioregional model. J Exp Mar Bio Ecol. 2007;350:320. [DOI]
    Ondiviela B, Losada IJ, Lara JL, Maza M, Galván C, Bouma TJ, et al. The role of seagrasses in coastal protection in a changing climate. Coast Eng. 2014;87:15868. [DOI]
    Rengasamy KRR, Sadeer NB, Zengin G, Mahomoodally MF, Cziáky Z, Jekő J, et al. Biopharmaceutical potential, chemical profile and in silico study of the seagrass– Syringodium isoetifolium (Asch.) Dandy. S Afr J Bot. 2019;127:16775. [DOI]
    McKenzie LJ, Nordlund LM, Jones BL, Cullen-Unsworth LC, Roelfsema C, Unsworth RKF. The global distribution of seagrass meadows. Environ Res Lett. 2020;15:074041. [DOI]
    Henrickson SE, Wong T, Allen P, Ford T, Epstein PR. Marine swimming-related illness: implications for monitoring and environmental policy. Environ Health Perspect. 2001;109:64550. [DOI] [PubMed] [PMC]
    Lamb JB, van de Water JA, Bourne DG, Altier C, Hein MY, Fiorenza EA, et al. Seagrass ecosystems reduce exposure to bacterial pathogens of humans, fishes, and invertebrates. Science. 2017;355:7313. [DOI] [PubMed]
    Ascioti FA, Mangano MC, Marcianò C, Sarà G. The sanitation service of seagrasses – dependencies and implications for the estimation of avoided costs. Ecosyst Serv. 2022;54:101418. [DOI]
    Zhao L, Ru S, He J, Zhang Z, Song X, Wang D, et al. Eelgrass (Zostera marina) and its epiphytic bacteria facilitate the sinking of microplastics in the seawater. Environ Pollut. 2022;292:118337. [DOI] [PubMed]
    Reusch TBH, Schubert PR, Marten SM, Gill D, Karez R, Busch K, et al. Lower Vibrio spp. abundances in Zostera marina leaf canopies suggest a novel ecosystem function for temperate seagrass beds. Mar Biol. 2021;168:149. [DOI]
    Alsaffar Z, Pearman JK, Cúrdia J, Ellis J, Calleja ML, Ruiz-Compean P, et al. The role of seagrass vegetation and local environmental conditions in shaping benthic bacterial and macroinvertebrate communities in a tropical coastal lagoon. Sci Rep. 2020;10:13550. [DOI] [PubMed] [PMC]
    Tarquinio F, Hyndes GA, Laverock B, Koenders A, Säwström C. The seagrass holobiont: understanding seagrass-bacteria interactions and their role in seagrass ecosystem functioning. FEMS Microbiol Lett. 2019;366:fnz057. [DOI] [PubMed]
    Conte C, Rotini A, Manfra L, D’Andrea MM, Winters G, Migliore L. The seagrass holobiont: What we know and what we still need to disclose for its possible use as an ecological indicator. Water. 2021;13:406. [DOI]
    Mohr W, Lehnen N, Ahmerkamp S, Marchant HK, Graf JS, Tschitschko B, et al. Terrestrial-type nitrogen-fixing symbiosis between seagrass and a marine bacterium. Nature. 2021;600:1059. [DOI] [PubMed] [PMC]
    Iqbal MM, Nishimura M, Haider MN, Sano M, Ijichi M, Kogure K, et al. Diversity and composition of microbial communities in an eelgrass (Zostera marina) bed in Tokyo Bay, Japan. Microbes Environ. 2021;36:ME21037. [DOI] [PubMed] [PMC]
    Trevathan-Tackett SM, Lane AL, Bishop N, Ross C. Metabolites derived from the tropical seagrass Thalassia testudinum are bioactive against pathogenic Labyrinthula sp. Aquat Bot. 2015;122:18. [DOI]
    Tarquinio F, Attlan O, Vanderklift MA, Berry O, Bissett A. Distinct endophytic bacterial communities inhabiting seagrass seeds. Front Microbiol. 2021;12:703014. [DOI] [PubMed] [PMC]
    Hurtado-McCormick V, Kahlke T, Petrou K, Jeffries T, Ralph PJ, Seymour JR. Regional and microenvironmental scale characterization of the Zostera muelleri seagrass Microbiome. Front Microbiol. 2019;10:1011. Erratum in: Front Microbiol. 2021;12:642964. [DOI] [PubMed] [PMC]
    Conte C, Apostolaki ET, Vizzini S, Migliore L. A tight interaction between the native seagrass Cymodocea nodosa and the exotic Halophila stipulacea in the Aegean Sea highlights seagrass holobiont variations. Plants (Basel). 2023;12:350. [DOI] [PubMed] [PMC]
    Mohapatra M, Manu S, Dash SP, Rastogi G. Seagrasses and local environment control the bacterial community structure and carbon substrate utilization in brackish sediments. J Environ Manage. 2022;314:115013. [DOI] [PubMed]
    Banister RB, Schwarz MT, Fine M, Ritchie KB, Muller EM. Instability and stasis among the microbiome of seagrass leaves, roots and rhizomes, and nearby sediments within a natural pH gradient. Microb Ecol. 2022;84:70316. Erratum in: Microb Ecol. 2023;85:1634. [DOI] [PubMed] [PMC]
    Vasarri M, De Biasi AM, Barletta E, Pretti C, Degl’Innocenti D. An overview of new insights into the benefits of the seagrass Posidonia oceanica for human health. Mar Drugs. 2021;19:476. [DOI] [PubMed] [PMC]
    Terrados J, Borum J. Why are seagrasses important? - Goods and services provided by seagrass meadows. In: Borum J, Duarte CM, Krause-Jensen D, Greve TM, editors. European seagrasses: an introduction to monitoring and management. The M&MS project; 2004. pp. 8–10.
    Zidorn C. Secondary metabolites of seagrasses (Alismatales and Potamogetonales; Alismatidae): chemical diversity, bioactivity, and ecological function. Phytochemistry. 2016;124:528. [DOI] [PubMed]
    Jiang Y, Li Q, Chen X, Jiang C. Isolation and cultivation methods of actinobacteria. In: Dhanasekaran D, Jiang Y, editors. Actinobacteria. Rijeka: IntechOpen; 2016. [DOI]
    Wu H, Chen W, Wang G, Dai S, Zhou D, Zhao H, et al. Culture-dependent diversity of actinobacteria associated with seagrass (Thalassia hemprichii). Afr J Microbiol Res. 2012;6:8794. [DOI]
    Ravikumar S, Gnanadesigan M, Saravanan A, Monisha N, Brindha V, Muthumari S. Antagonistic properties of seagrass associated Streptomyces sp. RAUACT-1: a source for anthraquinone rich compound. Asian Pac J Trop Med. 2012;5:88790. [DOI] [PubMed]
    Jose PA, sundari IS, Sivakala KK, Jebakumar SRD. Molecular phylogeny and plant growth promoting traits of endophytic bacteria isolated from roots of seagrass Cymodocea serrulata. Indian J Geo-Mar Sci. 2014;43:5719.
    Bibi F, Naseer MI, Hassan AM, Yasir M, Al-Ghamdi AAK, Azhar EI. Diversity and antagonistic potential of bacteria isolated from marine grass Halodule uninervis. 3 Biotech. 2018;8:48. [DOI] [PubMed] [PMC]
    Ettinger CL, Eisen JA. Fungi, bacteria and oomycota opportunistically isolated from the seagrass, Zostera marina. PLoS One. 2020;15:e0236135. Erratum in: PLoS One. 2021;16:e0251536. [DOI] [PubMed] [PMC]
    Cristianawati O, Sibero MT, Ayuningrum D, Nuryadi H, Syafitri E, Riniarsih I, et al. Screening of antibacterial activity of seagrass-associated bacteria from the North Java Sea, Indonesia against multidrug-resistant bacteria. Aquac Aquarium Conserv Legis. 2019;12:105464.
    Hamdy AHA, El-Fiky NM, El-Beih AA, Mohammed MM, Mettwally WS. Egyptian red sea seagrass as a source of biologically active secondary metabolites. Egypt Pharm J. 2020;19:22437. [DOI]
    Ragupathi Raja Kannan R, Arumugam R, Anantharaman P. Chemical composition and antibacterial activity of Indian seagrasses against urinary tract pathogens. Food Chem. 2012;135:24703. [DOI] [PubMed]
    Engel S, Puglisi MP, Jensen PR, Fenical W. Antimicrobial activities of extracts from tropical Atlantic marine plants against marine pathogens and saprophytes. Mar Biol. 2006;149:9911002. [DOI]
    Kim DH, Mahomoodally MF, Sadeer NB, Seok PG, Zengin G, Palaniveloo K, et al. Nutritional and bioactive potential of seagrasses: a review. S Afr J Bot. 2021;137:21627. [DOI]
    Gono CMP, Ahmadi P, Hertiani T, Septiana E, Putra MY, Chianese G. A comprehensive update on the bioactive compounds from seagrasses. Mar Drugs. 2022;20:406. [DOI] [PubMed] [PMC]
    Jafriati J, Hatta M, Yuniar N, Ade RJ, Dwiyanti R, Sabir M, et al. Thalassia hemprichii seagrass extract as antimicrobial and antioxidant potential on human: a mini review of the benefits of seagrass. J Biol Sci. 2019;19:36371. [DOI]
    Nur RM, Nurafni, Koroy K, Alwi D, Wahab I, Sulistiawati S, et al. The antibacterial activity of seagrass Enhalus acoroides against Staphylococcus aureus. IOP Conf Ser: Earth Environ Sci. 2021;890:012013. [DOI]
    De Vincenti L, Glasenapp Y, Cattò C, Villa F, Cappitelli F, Papenbrock J. Hindering the formation and promoting the dispersion of medical biofilms: non-lethal effects of seagrass extracts. BMC Complement Altern Med. 2018;18:168. [DOI] [PubMed] [PMC]
    Sun Y, Song Z, Zhang H, Liu P, Hu X. Seagrass vegetation affect the vertical organization of microbial communities in sediment. Mar Environ Res. 2020;162:105174. [DOI] [PubMed]
    Crump BC, Wojahn JM, Tomas F, Mueller RS. Metatranscriptomics and amplicon sequencing reveal mutualisms in seagrass microbiomes. Front Microbiol. 2018;9:388. [DOI] [PubMed] [PMC]
    Rotini A, Conte C, Seveso D, Montano S, Galli P, Vai M, et al. Daily variation of the associated microbial community and the Hsp60 expression in the Maldivian seagrass Thalassia hemprichii. J Sea Res. 2020;156:101835. [DOI]
    Ugarelli K, Laas P, Stingl U. The microbial communities of leaves and roots associated with turtle grass (Thalassia testudinum) and manatee grass (Syringodium filliforme) are distinct from seawater and sediment communities, but are similar between species and sampling sites. Microorganisms. 2018;7:4. [DOI] [PubMed] [PMC]
    Jensen SI, Kühl M, Priemé A. Different bacterial communities associated with the roots and bulk sediment of the seagrass Zostera marina. FEMS Microbiol Ecol. 2007;62:10817. [DOI] [PubMed]
    Damayanti V, Rachma RN, Santoso I, Yasman Y, Maryanto AE. Fermentation of antimicrobial substances of Streptomyces sp. BCy isolated from seagrass Cymodocearotundata using two different media. AIP Conf Proc. 2018;2023:020143. [DOI]
    Almaary KS, Alharbi NS, Kadaikunnan S, Khaled JM, Rajivgandhi G, Ramachandran G, et al. Anti-bacterial effect of marine sea grasses mediated endophytic actinomycetes against K. pneumoniae. J King Saud Univ Sci. 2021;33:101528. [DOI]
    Inaba N, Trainer VL, Onishi Y, Ishii KI, Wyllie-Echeverria S, Imai I. Algicidal and growth-inhibiting bacteria associated with seagrass and macroalgae beds in Puget Sound, WA, USA. Harmful Algae. 2017;62:13647. [DOI] [PubMed]
    Martin BC, Gleeson D, Statton J, Siebers AR, Grierson P, Ryan MH, et al. Low light availability alters root exudation and reduces putative beneficial microorganisms in seagrass roots. Front Microbiol. 2018;8:2667. [DOI] [PubMed] [PMC]