﻿<?xml version="1.0" encoding="utf-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD JATS (Z39.96) Journal Publishing DTD v1.1 20151215//EN" "JATS-journalpublishing1.dtd">
<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" article-type="review-article">
<front>
<journal-meta>
<journal-id journal-id-type="nlm-ta">Explor Drug Sci</journal-id>
<journal-id journal-id-type="publisher-id">EDS</journal-id>
<journal-title-group>
<journal-title>Exploration of Drug Science</journal-title>
</journal-title-group>
<issn pub-type="epub">2836-7677</issn>
<publisher>
<publisher-name>Open Exploration Publishing</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.37349/eds.2024.00038</article-id>
<article-id pub-id-type="manuscript">100838</article-id>
<article-categories>
<subj-group>
<subject>Mini Review</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Mini-review on the antimicrobial potential of actinobacteria associated with seagrasses</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-5802-2777</contrib-id>
<name>
<surname>Siro</surname>
<given-names>Galana</given-names>
</name>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/">Writing—original draft</role>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing—review &amp; editing</role>
<xref ref-type="aff" rid="I1">
<sup>1</sup>
</xref>
<xref ref-type="corresp" rid="cor1">
<sup>*</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-4562-9852</contrib-id>
<name>
<surname>Pipite</surname>
<given-names>Atanas</given-names>
</name>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/">Writing—original draft</role>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing—review &amp; editing</role>
<role content-type="https://credit.niso.org/contributor-roles/supervision/">Supervision</role>
<xref ref-type="aff" rid="I2">
<sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="editor">
<name>
<surname>Reichardt</surname>
<given-names>Juergen</given-names>
</name>
<role>Academic Editor</role>
<aff>James Cook University, Australia</aff>
</contrib>
</contrib-group>
<aff id="I1">
<sup>1</sup>School of Agriculture, Geography, Environment, Ocean and Natural Sciences (SAGEONS), Discipline of Biological and Chemical Sciences, The University of the South Pacific, Laucala Campus, Suva 1168, Fiji</aff>
<aff id="I2">
<sup>2</sup>School of Natural Sciences, Department of Biomolecular, Massey University, Palmerston North 4410, New Zealand</aff>
<author-notes>
<corresp id="cor1">
<sup>*</sup>
<bold>Correspondence:</bold> Galana Siro, School of Agriculture, Geography, Environment, Ocean and Natural Sciences (SAGEONS), Discipline of Biological and Chemical Sciences, The University of the South Pacific, Laucala Campus, Suva 1168, Fiji. <email>sirogalana@yahoo.com</email></corresp>
</author-notes>
<pub-date pub-type="ppub">
<year>2024</year>
</pub-date>
<pub-date pub-type="epub">
<day>29</day>
<month>02</month>
<year>2024</year>
</pub-date>
<volume>2</volume>
<issue>1</issue>
<fpage>117</fpage>
<lpage>125</lpage>
<history>
<date date-type="received">
<day>10</day>
<month>08</month>
<year>2023</year>
</date>
<date date-type="accepted">
<day>01</day>
<month>12</month>
<year>2023</year>
</date>
</history>
<permissions>
<copyright-statement>© The Author(s) 2024.</copyright-statement>
<license xlink:href="https://creativecommons.org/licenses/by/4.0/">
<license-p>This is an Open Access article licensed under a Creative Commons Attribution 4.0 International License (<ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">https://creativecommons.org/licenses/by/4.0/</ext-link>), which permits unrestricted use, sharing, adaptation, distribution and reproduction in any medium or format, for any purpose, even commercially, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made.</license-p>
</license>
</permissions>
<abstract>
<p>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.</p>
</abstract>
<kwd-group>
<kwd>Actinobacteria</kwd>
<kwd>seagrasses</kwd>
<kwd>biosynthetic metabolites</kwd>
<kwd>marine ecosystem</kwd>
<kwd>antimicrobial resistance</kwd>
</kwd-group>
</article-meta>
</front>
<body>
<sec id="s1">
<title>Introduction</title>
<p id="p-1">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 [<xref ref-type="bibr" rid="B1">1</xref>, <xref ref-type="bibr" rid="B2">2</xref>]. As a counter approach, the administration of antibiotics and the tedious bioprospects of novel bioactive compounds from marine sources are common [<xref ref-type="bibr" rid="B3">3</xref>, <xref ref-type="bibr" rid="B4">4</xref>]. A quantum of information regarding the natural products from marine environments has received great recognition due to their diverse applications and efficiency [<xref ref-type="bibr" rid="B5">5</xref>]. Bioprospects in marine habitats have continuously provided a series of novel and interesting metabolites [<xref ref-type="bibr" rid="B6">6</xref>–<xref ref-type="bibr" rid="B8">8</xref>]. An important marine ecosystem represented in this review is seagrasses. Seagrass meadows support gradation in species diversity and high biodiversity of marine life [<xref ref-type="bibr" rid="B9">9</xref>]. Actinobacteria sometimes referred to as actinomycetes are Gram-positive bacteria and form a unique group of bacterial lineages. They belong to the phylum Actinomycetota (<uri xlink:href="https://lpsn.dsmz.de/phylum/actinomycetota">https://lpsn.dsmz.de/phylum/actinomycetota</uri>) and are regarded as hotspots for clinical drug discovery. Their genomes have a high guanine-cytosine (GC) content and genes associated with antibiotic production [<xref ref-type="bibr" rid="B10">10</xref>–<xref ref-type="bibr" rid="B12">12</xref>]. The genus <italic>Streptomyces</italic> is the most widely gifted group of bacteria with efficient antibiotic machinery and is responsible for the high availability of clinical antibiotics [<xref ref-type="bibr" rid="B13">13</xref>, <xref ref-type="bibr" rid="B14">14</xref>]. 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 [<xref ref-type="bibr" rid="B15">15</xref>, <xref ref-type="bibr" rid="B16">16</xref>]. 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 [<xref ref-type="bibr" rid="B17">17</xref>–<xref ref-type="bibr" rid="B19">19</xref>]. 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.</p>
</sec>
<sec id="s2">
<title>Seagrass biology, evolution, and biodiversity</title>
<p id="p-2">Seagrasses are submerged, marine flowering plants that thrive in the ocean except in the polar regions [<xref ref-type="bibr" rid="B20">20</xref>]. They comprise over 60 species, belonging to four exclusive plant families including Zosteraceae, Hydrocharitaceae, Posidoniaceae, and Cymodoceaceae, and support immense levels of biodiversity [<xref ref-type="bibr" rid="B21">21</xref>–<xref ref-type="bibr" rid="B25">25</xref>]. Seagrasses have a range of features that arose from adapting to living in submerged marine conditions [<xref ref-type="bibr" rid="B26">26</xref>, <xref ref-type="bibr" rid="B27">27</xref>]. As an aquatic angiosperm, their multiple colonization events in the ocean predated over 70 million years ago [<xref ref-type="bibr" rid="B25">25</xref>]. Notably, seagrasses are rhizomatous plants adapted to aquatic environments [<xref ref-type="bibr" rid="B28">28</xref>]. Depending on their growth sizes, seagrasses range from small species (e.g., <italic>Halophila engelmannii</italic>) to large species (e.g., <italic>Posidonia australis</italic>) [<xref ref-type="bibr" rid="B23">23</xref>]. The highly productive nature of seagrasses is largely influenced by a mixture of sexual and asexual clonal reproduction [<xref ref-type="bibr" rid="B26">26</xref>]. 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) [<xref ref-type="bibr" rid="B24">24</xref>, <xref ref-type="bibr" rid="B28">28</xref>, <xref ref-type="bibr" rid="B29">29</xref>]. The ecological importance of seagrass meadows is irrefutable, and yet remains poorly studied, in terms of medicinal properties [<xref ref-type="bibr" rid="B30">30</xref>]. 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 [<xref ref-type="bibr" rid="B31">31</xref>]. 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 [<xref ref-type="bibr" rid="B28">28</xref>]. Its mass coverage spans from small patches to extensive underwater lawns [<xref ref-type="bibr" rid="B28">28</xref>].</p>
<p id="p-3">The ocean often receives a cocktail of raw sewage or pollutants. Consequently, shoreline microbial populations can spike to dangerous levels with serious health implications [<xref ref-type="bibr" rid="B32">32</xref>]. 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 [<xref ref-type="bibr" rid="B33">33</xref>–<xref ref-type="bibr" rid="B36">36</xref>]. 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) [<xref ref-type="bibr" rid="B37">37</xref>]. 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 [<xref ref-type="bibr" rid="B26">26</xref>, <xref ref-type="bibr" rid="B38">38</xref>–<xref ref-type="bibr" rid="B41">41</xref>]. Both the seagrass host and their associate microbes are capable of producing potent metabolites that prevent the invasion of opportunistic pathogens [<xref ref-type="bibr" rid="B39">39</xref>, <xref ref-type="bibr" rid="B42">42</xref>]. Studies have shown that distinct microbial communities exist in discrete microenvironments of seagrasses [<xref ref-type="bibr" rid="B43">43</xref>]. 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 [<xref ref-type="bibr" rid="B44">44</xref>–<xref ref-type="bibr" rid="B47">47</xref>].</p>
</sec>
<sec id="s3">
<title>Traditional and medicinal use of seagrass around the world</title>
<p id="p-4">There is evidence throughout history of the use of seagrass as food, medicine, fertilizer, and livestock feed [<xref ref-type="bibr" rid="B20">20</xref>, <xref ref-type="bibr" rid="B48">48</xref>, <xref ref-type="bibr" rid="B49">49</xref>]. 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) [<xref ref-type="bibr" rid="B20">20</xref>]. 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) [<xref ref-type="bibr" rid="B20">20</xref>]. In Tunisia, the leaves of the seagrass, <italic>Posidonia oceanica</italic>, which have antifungal and insect repellent properties, have been used as livestock bedding and as feed supplements for poultry and livestock [<xref ref-type="bibr" rid="B48">48</xref>]. 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) [<xref ref-type="bibr" rid="B49">49</xref>].</p>
<p id="p-5">The use of seagrass throughout history has shown that there is knowledge of the wide range of qualities possessed by these oceanic plants [<xref ref-type="bibr" rid="B50">50</xref>]. 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 [<xref ref-type="bibr" rid="B50">50</xref>]. Remarkably, a total of 154 natural compounds derived from 70 seagrass species have been reported so far, predominantly from the host seagrass itself [<xref ref-type="bibr" rid="B50">50</xref>].</p>
</sec>
<sec id="s4">
<title>Diversity and bioactive profiles of seagrass-derived actinobacteria</title>
<p id="p-6">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 [<xref ref-type="bibr" rid="B51">51</xref>]. 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 [<xref ref-type="bibr" rid="B52">52</xref>–<xref ref-type="bibr" rid="B57">57</xref>]. 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 <xref ref-type="table" rid="t1">Table 1</xref>.</p>
<table-wrap id="t1">
<label>Table 1</label>
<caption>
<p>Genus/species of actinobacteria isolated from seagrasses between 2012 and 2020</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th>
<bold>Genus/species</bold>
</th>
<th>
<bold>Family</bold>
</th>
<th>
<bold>Seagrass species and nature of sample</bold>
</th>
<th>
<bold>Bioactivity</bold>
</th>
<th>
<bold>Country</bold>
</th>
<th>
<bold>Reference</bold>
</th>
</tr>
</thead>
<tbody>
<tr>
<td>
<italic>Streptomyces</italic> spp., <italic>Micromonospora</italic> spp., <italic>Verrucosispora</italic> (<italic>Micromonospora</italic>) sp., <italic>Saccharomonospora</italic> spp., <italic>Actinomycetospora</italic> sp., <italic>Microbacterium</italic> sp., <italic>Mycobacterium</italic> spp., <italic>Nonomuraea</italic> sp., <italic>Nocardiopsis</italic> sp., and <italic>Glycomyces</italic> sp.</td>
<td>Streptomycetaceae, Micromonosporaceae, Pseudonocardiaceae, Microbacteriaceae, Mycobacteriaceae, Streptosporangiaceae, Nocardiopsidaceae, and Glycomycetaceae</td>
<td>
<italic>Thalassia hemprichii</italic> (whole plant)</td>
<td>Contain nonribosomal peptide synthetase (<italic>NRPS</italic>) and polyketide synthase (<italic>PKS</italic>) genes (antibacterial activity)</td>
<td>China</td>
<td>[<xref ref-type="bibr" rid="B52">52</xref>]</td>
</tr>
<tr>
<td>
<italic>Streptomyces</italic> spp.</td>
<td>Streptomycetaceae</td>
<td>
<italic>Syringodium isoetifolium</italic> (leaves and roots)</td>
<td>Antibacterial activity</td>
<td>India</td>
<td>[<xref ref-type="bibr" rid="B53">53</xref>]</td>
</tr>
<tr>
<td>
<italic>Saccharomonospora</italic> sp., <italic>Kocuria</italic> sp.</td>
<td>Pseudonocardiaceae, Micrococcaceae</td>
<td>
<italic>Cymodocea serrulate</italic> (roots)</td>
<td>Phosphate solubilizing, nitrogen-fixing ability, and enzyme activity</td>
<td>Kasuwari Island, India</td>
<td>[<xref ref-type="bibr" rid="B54">54</xref>]</td>
</tr>
<tr>
<td>
<italic>Arthrobacter</italic> spp.</td>
<td>Micrococcaceae</td>
<td>
<italic>Zostera marina</italic> and <italic>Zostera japonica</italic> (leaves)</td>
<td>Algicidal activity</td>
<td>Puget Sound, USA</td>
<td>[<xref ref-type="bibr" rid="B73">73</xref>]</td>
</tr>
<tr>
<td>
<italic>Kocuria palustris</italic>, <italic>Kocuria atrinae</italic>, <italic>Arthrobacter flavus</italic>, <italic>Ornithinimicrobium humiphilum</italic>, <italic>Corynebacterium afermentans</italic> subsp. <italic>afermentans</italic></td>
<td>Micrococcaceae, Ornithinimicrobiaceae, and Corynebacteriaceae</td>
<td>
<italic>Halodule uninervis</italic> (soil and root)</td>
<td>Antifungal activity</td>
<td>Saudi Arabia</td>
<td>[<xref ref-type="bibr" rid="B55">55</xref>]</td>
</tr>
<tr>
<td>Not available</td>
<td>Cellulomonadaceae, Microbacteriaceae</td>
<td>
<italic>Halophila ovalis</italic> (roots)</td>
<td>No bioactive test performed</td>
<td>Australia</td>
<td>[<xref ref-type="bibr" rid="B74">74</xref>]</td>
</tr>
<tr>
<td>
<italic>Streptomyces</italic> sp.</td>
<td>Streptomycetaceae</td>
<td>
<italic>Cymodocea rotundata</italic> (whole plant)</td>
<td>Antimicrobial activity</td>
<td>Indonesia</td>
<td>[<xref ref-type="bibr" rid="B71">71</xref>]</td>
</tr>
<tr>
<td>
<italic>Streptomyces lienomycini</italic>
</td>
<td>Streptomycetaceae</td>
<td>
<italic>Enhalus acoroides</italic> (leaves)</td>
<td>Antibacterial activity</td>
<td>Indonesia</td>
<td>[<xref ref-type="bibr" rid="B57">57</xref>]</td>
</tr>
<tr>
<td>
<italic>Isoptericola</italic> sp., <italic>Rhodococcus</italic> sp., and <italic>Streptomyces</italic> spp.</td>
<td>Promicromonosporaceae, Nocardiaceae, and Streptomycetaceae</td>
<td>
<italic>Zostera marina</italic> (leaf and associate sediment)</td>
<td>No bioactive test performed</td>
<td>Bodega Bay, USA</td>
<td>[<xref ref-type="bibr" rid="B56">56</xref>]</td>
</tr>
</tbody>
</table>
</table-wrap>
<p id="p-7">Seagrasses harbor a rich pool of specialized metabolites [<xref ref-type="bibr" rid="B58">58</xref>, <xref ref-type="bibr" rid="B59">59</xref>]. 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 [<xref ref-type="bibr" rid="B25">25</xref>, <xref ref-type="bibr" rid="B60">60</xref>–<xref ref-type="bibr" rid="B65">65</xref>]. 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 [<xref ref-type="bibr" rid="B66">66</xref>]. 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 [<xref ref-type="bibr" rid="B9">9</xref>, <xref ref-type="bibr" rid="B67">67</xref>–<xref ref-type="bibr" rid="B69">69</xref>]. Actinobacteria are among the pioneer colonizers of seagrasses [<xref ref-type="bibr" rid="B70">70</xref>] and their diversity varies between seagrass species [<xref ref-type="bibr" rid="B52">52</xref>, <xref ref-type="bibr" rid="B53">53</xref>]. The seagrass-derived actinomycetes are capable of producing antimicrobial agents (<xref ref-type="table" rid="t1">Table 1</xref>). 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 [<xref ref-type="bibr" rid="B52">52</xref>, <xref ref-type="bibr" rid="B53">53</xref>, <xref ref-type="bibr" rid="B57">57</xref>, <xref ref-type="bibr" rid="B71">71</xref>, <xref ref-type="bibr" rid="B72">72</xref>], antifungal activity [<xref ref-type="bibr" rid="B55">55</xref>, <xref ref-type="bibr" rid="B71">71</xref>], algicidal activity [<xref ref-type="bibr" rid="B73">73</xref>], nitrogen-fixing ability, and enzymatic activity [<xref ref-type="bibr" rid="B54">54</xref>]. Conspicuously, there are available studies where no bioactive test was performed for the actinobacteria discovered [<xref ref-type="bibr" rid="B56">56</xref>, <xref ref-type="bibr" rid="B74">74</xref>]. Interestingly, Wu et al. [<xref ref-type="bibr" rid="B52">52</xref>] reported the isolation of <italic>Verrucosispora</italic> sp. from the seagrass <italic>Thalassia hemprichii</italic> (this was the first time a <italic>Verrucosispora</italic> strain was isolated as a plant endophyte). The members of the genus <italic>Verrucosispora</italic> have subsequently all been transferred to the genus <italic>Micromonospora</italic> (<uri xlink:href="https://lpsn.dsmz.de/genus/verrucosispora">https://lpsn.dsmz.de/genus/verrucosispora</uri>).</p>
<p id="p-8">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.</p>
</sec>
<sec id="s5">
<title>Conclusions</title>
<p id="p-9">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 [<xref ref-type="bibr" rid="B50">50</xref>, <xref ref-type="bibr" rid="B58">58</xref>]. 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.</p>
</sec>
</body>
<back>
<sec id="s6">
<title>Declarations</title>
<sec>
<title>Author contributions</title>
<p>GS: Conceptualization, Writing—original draft, Writing—review &amp; editing. AP: Conceptualization, Writing—original draft, Writing—review &amp; editing, Supervision. All authors read and approved the submitted version.</p>
</sec>
<sec sec-type="COI-statement">
<title>Conflicts of interest</title>
<p>The authors declare that they have no conflicts of interest.</p>
</sec>
<sec>
<title>Ethical approval</title>
<p>Not applicable.</p>
</sec>
<sec>
<title>Consent to participate</title>
<p>Not applicable.</p>
</sec>
<sec>
<title>Consent to publication</title>
<p>Not applicable.</p>
</sec>
<sec sec-type="data-availability">
<title>Availability of data and materials</title>
<p>Not applicable.</p>
</sec>
<sec>
<title>Funding</title>
<p>Not applicable.</p>
</sec>
<sec>
<title>Copyright</title>
<p>© The Author(s) 2024.</p>
</sec>
</sec>
<ref-list>
<ref id="B1">
<label>1</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Munk</surname>
<given-names>P</given-names>
</name>
<name>
<surname>Brinch</surname>
<given-names>C</given-names>
</name>
<name>
<surname>Møller</surname>
<given-names>FD</given-names>
</name>
<name>
<surname>Petersen</surname>
<given-names>TN</given-names>
</name>
<name>
<surname>Hendriksen</surname>
<given-names>RS</given-names>
</name>
<name>
<surname>Seyfarth</surname>
<given-names>AM</given-names>
</name>
<etal>et al.</etal>
<collab>Global Sewage Surveillance Consortium</collab>
<name>
<surname>Larsson</surname>
<given-names>DGJ</given-names>
</name>
<name>
<surname>Koopmans</surname>
<given-names>M</given-names>
</name>
<name>
<surname>Woolhouse</surname>
<given-names>M</given-names>
</name>
<name>
<surname>Aarestrup</surname>
<given-names>FM</given-names>
</name>
</person-group>
<article-title>Genomic analysis of sewage from 101 countries reveals global landscape of antimicrobial resistance</article-title>
<source>Nat Commun</source>
<year iso-8601-date="2022">2022</year>
<volume>13</volume>
<elocation-id>7251</elocation-id>
<comment>Erratum in: Nat Commun. 2023;14:178.</comment>
<pub-id pub-id-type="doi">10.1038/s41467-022-34312-7</pub-id><pub-id pub-id-type="pmid">36456547</pub-id><pub-id pub-id-type="pmcid">PMC9715550</pub-id></element-citation>
</ref>
<ref id="B2">
<label>2</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Brinch</surname>
<given-names>C</given-names>
</name>
<name>
<surname>Leekitcharoenphon</surname>
<given-names>P</given-names>
</name>
<name>
<surname>Duarte</surname>
<given-names>ASR</given-names>
</name>
<name>
<surname>Svendsen</surname>
<given-names>CA</given-names>
</name>
<name>
<surname>Jensen</surname>
<given-names>JD</given-names>
</name>
<name>
<surname>Aarestrup</surname>
<given-names>FM</given-names>
</name>
</person-group>
<article-title>Long-term temporal stability of the resistome in sewage from Copenhagen</article-title>
<source>mSystems</source>
<year iso-8601-date="2020">2020</year>
<volume>5</volume>
<elocation-id>e00841-20</elocation-id>
<pub-id pub-id-type="doi">10.1128/mSystems.00841-20</pub-id><pub-id pub-id-type="pmid">33082278</pub-id><pub-id pub-id-type="pmcid">PMC7577296</pub-id></element-citation>
</ref>
<ref id="B3">
<label>3</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Voser</surname>
<given-names>TM</given-names>
</name>
<name>
<surname>Campbell</surname>
<given-names>MD</given-names>
</name>
<name>
<surname>Carroll</surname>
<given-names>AR</given-names>
</name>
</person-group>
<article-title>How different are marine microbial natural products compared to their terrestrial counterparts?</article-title>
<source>Nat Prod Rep</source>
<year iso-8601-date="2022">2022</year>
<volume>39</volume>
<fpage>7</fpage>
<lpage>19</lpage>
<pub-id pub-id-type="doi">10.1039/d1np00051a</pub-id><pub-id pub-id-type="pmid">34651634</pub-id></element-citation>
</ref>
<ref id="B4">
<label>4</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Sigwart</surname>
<given-names>JD</given-names>
</name>
<name>
<surname>Blasiak</surname>
<given-names>R</given-names>
</name>
<name>
<surname>Jaspars</surname>
<given-names>M</given-names>
</name>
<name>
<surname>Jouffray</surname>
<given-names>JB</given-names>
</name>
<name>
<surname>Tasdemir</surname>
<given-names>D</given-names>
</name>
</person-group>
<article-title>Unlocking the potential of marine biodiscovery</article-title>
<source>Nat Prod Rep</source>
<year iso-8601-date="2021">2021</year>
<volume>38</volume>
<fpage>1235</fpage>
<lpage>42</lpage>
<pub-id pub-id-type="doi">10.1039/d0np00067a</pub-id><pub-id pub-id-type="pmid">34287433</pub-id></element-citation>
</ref>
<ref id="B5">
<label>5</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Carroll</surname>
<given-names>AR</given-names>
</name>
<name>
<surname>Copp</surname>
<given-names>BR</given-names>
</name>
<name>
<surname>Davis</surname>
<given-names>RA</given-names>
</name>
<name>
<surname>Keyzers</surname>
<given-names>RA</given-names>
</name>
<name>
<surname>Prinsep</surname>
<given-names>MR</given-names>
</name>
</person-group>
<article-title>Marine natural products</article-title>
<source>Nat Prod Rep</source>
<year iso-8601-date="2022">2022</year>
<volume>39</volume>
<fpage>1122</fpage>
<lpage>71</lpage>
<pub-id pub-id-type="doi">10.3390/md20080528</pub-id><pub-id pub-id-type="pmid">36005531</pub-id><pub-id pub-id-type="pmcid">PMC9410185</pub-id></element-citation>
</ref>
<ref id="B6">
<label>6</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Siro</surname>
<given-names>G</given-names>
</name>
<name>
<surname>Pipite</surname>
<given-names>A</given-names>
</name>
<name>
<surname>Christi</surname>
<given-names>K</given-names>
</name>
<name>
<surname>Srinivasan</surname>
<given-names>S</given-names>
</name>
<name>
<surname>Subramani</surname>
<given-names>R</given-names>
</name>
</person-group>
<article-title>Marine actinomycetes associated with stony corals: a potential hotspot for specialized metabolites</article-title>
<source>Microorganisms</source>
<year iso-8601-date="2022">2022</year>
<volume>10</volume>
<elocation-id>1349</elocation-id>
<pub-id pub-id-type="doi">10.3390/microorganisms10071349</pub-id><pub-id pub-id-type="pmid">35889068</pub-id><pub-id pub-id-type="pmcid">PMC9319285</pub-id></element-citation>
</ref>
<ref id="B7">
<label>7</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Siro</surname>
<given-names>G</given-names>
</name>
<name>
<surname>Donald</surname>
<given-names>L</given-names>
</name>
<name>
<surname>Pipite</surname>
<given-names>A</given-names>
</name>
</person-group>
<article-title>The diversity of deep-sea actinobacteria and their natural products: an epitome of curiosity and drug discovery</article-title>
<source>Diversity</source>
<year iso-8601-date="2023">2023</year>
<volume>15</volume>
<elocation-id>30</elocation-id>
<pub-id pub-id-type="doi">10.3390/d15010030</pub-id></element-citation>
</ref>
<ref id="B8">
<label>8</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Khan</surname>
<given-names>N</given-names>
</name>
<name>
<surname>Yılmaz</surname>
<given-names>S</given-names>
</name>
<name>
<surname>Aksoy</surname>
<given-names>S</given-names>
</name>
<name>
<surname>Uzel</surname>
<given-names>A</given-names>
</name>
<name>
<surname>Tosun</surname>
<given-names>Ç</given-names>
</name>
<name>
<surname>Kirmizibayrak</surname>
<given-names>PB</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>Polyethers isolated from the marine actinobacterium <italic>Streptomyces cacaoi</italic> inhibit autophagy and induce apoptosis in cancer cells</article-title>
<source>Chem Biol Interact</source>
<year iso-8601-date="2019">2019</year>
<volume>307</volume>
<fpage>167</fpage>
<lpage>78</lpage>
<pub-id pub-id-type="doi">10.1016/j.cbi.2019.04.035</pub-id><pub-id pub-id-type="pmid">31059704</pub-id></element-citation>
</ref>
<ref id="B9">
<label>9</label>
<element-citation publication-type="book">
<person-group person-group-type="author">
<name>
<surname>Borowitzka</surname>
<given-names>MA</given-names>
</name>
<name>
<surname>Lavery</surname>
<given-names>PS</given-names>
</name>
<name>
<surname>Keulen</surname>
<given-names>M</given-names>
</name>
</person-group>
<article-title>Epiphytes of seagrasses</article-title>
<person-group person-group-type="editor">
<name>
<surname>Larkum</surname>
<given-names>AWD</given-names>
</name>
<name>
<surname>Orth</surname>
<given-names>RJ</given-names>
</name>
<name>
<surname>Duarte</surname>
<given-names>CM</given-names>
</name>
</person-group>
<source>Seagrasses: biology, ecology and conservation</source>
<publisher-loc>Dordrecht</publisher-loc>
<publisher-name>Springer Netherlands</publisher-name>
<year iso-8601-date="2006">2006</year>
<comment>pp. 441–61.</comment>
</element-citation>
</ref>
<ref id="B10">
<label>10</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Mast</surname>
<given-names>Y</given-names>
</name>
<name>
<surname>Stegmann</surname>
<given-names>E</given-names>
</name>
</person-group>
<article-title>Actinomycetes: the antibiotics producers</article-title>
<source>Antibiotics</source>
<year iso-8601-date="2019">2019</year>
<volume>8</volume>
<elocation-id>105</elocation-id>
<pub-id pub-id-type="doi">10.3390/antibiotics8030105</pub-id></element-citation>
</ref>
<ref id="B11">
<label>11</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Barka</surname>
<given-names>EA</given-names>
</name>
<name>
<surname>Vatsa</surname>
<given-names>P</given-names>
</name>
<name>
<surname>Sanchez</surname>
<given-names>L</given-names>
</name>
<name>
<surname>Gaveau-Vaillant</surname>
<given-names>N</given-names>
</name>
<name>
<surname>Jacquard</surname>
<given-names>C</given-names>
</name>
<name>
<surname>Meier-Kolthoff</surname>
<given-names>JP</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>Taxonomy, physiology, and natural products of actinobacteria</article-title>
<source>Microbiol Mol Biol Rev</source>
<year iso-8601-date="2015">2015</year>
<volume>80</volume>
<fpage>1</fpage>
<lpage>43</lpage>
<comment>Erratum in: Microbiol Mol Biol Rev. 2016;80:iii.</comment>
<pub-id pub-id-type="doi">10.1128/MMBR.00019-15</pub-id><pub-id pub-id-type="pmid">26609051</pub-id><pub-id pub-id-type="pmcid">PMC4711186</pub-id></element-citation>
</ref>
<ref id="B12">
<label>12</label>
<element-citation publication-type="book">
<person-group person-group-type="author">
<name>
<surname>Anandan</surname>
<given-names>R</given-names>
</name>
<name>
<surname>Dharumadurai</surname>
<given-names>D</given-names>
</name>
<name>
<surname>Manogaran</surname>
<given-names>GP</given-names>
</name>
</person-group>
<article-title>An introduction to actinobacteria</article-title>
<person-group person-group-type="editor">
<name>
<surname>Dhanasekaran</surname>
<given-names>D</given-names>
</name>
<name>
<surname>Jiang</surname>
<given-names>Y</given-names>
</name>
</person-group>
<source>Actinobacteria</source>
<publisher-loc>Rijeka</publisher-loc>
<publisher-name>IntechOpen</publisher-name>
<year iso-8601-date="2016">2016</year>
<pub-id pub-id-type="doi">10.5772/62329</pub-id></element-citation>
</ref>
<ref id="B13">
<label>13</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Watve</surname>
<given-names>MG</given-names>
</name>
<name>
<surname>Tickoo</surname>
<given-names>R</given-names>
</name>
<name>
<surname>Jog</surname>
<given-names>MM</given-names>
</name>
<name>
<surname>Bhole</surname>
<given-names>BD</given-names>
</name>
</person-group>
<article-title>How many antibiotics are produced by the genus <italic>Streptomyces</italic>?</article-title>
<source>Arch Microbiol</source>
<year iso-8601-date="2001">2001</year>
<volume>176</volume>
<fpage>386</fpage>
<lpage>90</lpage>
<pub-id pub-id-type="doi">10.1007/s002030100345</pub-id><pub-id pub-id-type="pmid">11702082</pub-id></element-citation>
</ref>
<ref id="B14">
<label>14</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Donald</surname>
<given-names>L</given-names>
</name>
<name>
<surname>Pipite</surname>
<given-names>A</given-names>
</name>
<name>
<surname>Subramani</surname>
<given-names>R</given-names>
</name>
<name>
<surname>Owen</surname>
<given-names>J</given-names>
</name>
<name>
<surname>Keyzers</surname>
<given-names>RA</given-names>
</name>
<name>
<surname>Taufa</surname>
<given-names>T</given-names>
</name>
</person-group>
<article-title>
<italic>Streptomyces</italic>: still the biggest producer of new natural secondary metabolites, a current perspective</article-title>
<source>Microbiol Res</source>
<year iso-8601-date="2022">2022</year>
<volume>13</volume>
<fpage>418</fpage>
<lpage>65</lpage>
<pub-id pub-id-type="doi">10.3390/microbiolres13030031</pub-id></element-citation>
</ref>
<ref id="B15">
<label>15</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Verma</surname>
<given-names>S</given-names>
</name>
<name>
<surname>Kuila</surname>
<given-names>A</given-names>
</name>
</person-group>
<article-title>Bioremediation of heavy metals by microbial process</article-title>
<source>Environ Technol Innov</source>
<year iso-8601-date="2019">2019</year>
<volume>14</volume>
<elocation-id>100369</elocation-id>
<pub-id pub-id-type="doi">10.1016/j.eti.2019.100369</pub-id></element-citation>
</ref>
<ref id="B16">
<label>16</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Sharma</surname>
<given-names>M</given-names>
</name>
<name>
<surname>Dangi</surname>
<given-names>P</given-names>
</name>
<name>
<surname>Choudhary</surname>
<given-names>M</given-names>
</name>
</person-group>
<article-title>Actinomycetes: source, identification, and their applications</article-title>
<source>Int J Curr Microbiol Appl Sci</source>
<year iso-8601-date="2014">2014</year>
<volume>3</volume>
<fpage>801</fpage>
<lpage>32</lpage>
</element-citation>
</ref>
<ref id="B17">
<label>17</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Singh</surname>
<given-names>R</given-names>
</name>
<name>
<surname>Dubey</surname>
<given-names>AK</given-names>
</name>
</person-group>
<article-title>Diversity and applications of endophytic actinobacteria of plants in special and other ecological niches</article-title>
<source>Front Microbiol</source>
<year iso-8601-date="2018">2018</year>
<volume>9</volume>
<elocation-id>1767</elocation-id>
<pub-id pub-id-type="doi">10.3389/fmicb.2018.01767</pub-id><pub-id pub-id-type="pmid">30135681</pub-id><pub-id pub-id-type="pmcid">PMC6092505</pub-id></element-citation>
</ref>
<ref id="B18">
<label>18</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Jagannathan</surname>
<given-names>SV</given-names>
</name>
<name>
<surname>Manemann</surname>
<given-names>EM</given-names>
</name>
<name>
<surname>Rowe</surname>
<given-names>SE</given-names>
</name>
<name>
<surname>Callender</surname>
<given-names>MC</given-names>
</name>
<name>
<surname>Soto</surname>
<given-names>W</given-names>
</name>
</person-group>
<article-title>Marine actinomycetes, new sources of biotechnological products</article-title>
<source>Mar Drugs</source>
<year iso-8601-date="2021">2021</year>
<volume>19</volume>
<elocation-id>365</elocation-id>
<pub-id pub-id-type="doi">10.3390/md19070365</pub-id><pub-id pub-id-type="pmid">34201951</pub-id><pub-id pub-id-type="pmcid">PMC8304352</pub-id></element-citation>
</ref>
<ref id="B19">
<label>19</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Lam</surname>
<given-names>KS</given-names>
</name>
</person-group>
<article-title>Discovery of novel metabolites from marine actinomycetes</article-title>
<source>Curr Opin Microbiol</source>
<year iso-8601-date="2006">2006</year>
<volume>9</volume>
<fpage>245</fpage>
<lpage>51</lpage>
<pub-id pub-id-type="doi">10.1016/j.mib.2006.03.004</pub-id><pub-id pub-id-type="pmid">16675289</pub-id></element-citation>
</ref>
<ref id="B20">
<label>20</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Rengasamy</surname>
<given-names>RRK</given-names>
</name>
<name>
<surname>Radjassegarin</surname>
<given-names>A</given-names>
</name>
<name>
<surname>Perumal</surname>
<given-names>A</given-names>
</name>
</person-group>
<article-title>Seagrasses as potential source of medicinal food ingredients: nutritional analysis and multivariate approach</article-title>
<source>Biomed Prev Nutr</source>
<year iso-8601-date="2013">2013</year>
<volume>3</volume>
<fpage>375</fpage>
<lpage>80</lpage>
<pub-id pub-id-type="doi">10.1016/j.bionut.2013.06.011</pub-id></element-citation>
</ref>
<ref id="B21">
<label>21</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Unsworth</surname>
<given-names>RKF</given-names>
</name>
<name>
<surname>Nordlund</surname>
<given-names>LM</given-names>
</name>
<name>
<surname>Cullen-Unsworth</surname>
<given-names>LC</given-names>
</name>
</person-group>
<article-title>Seagrass meadows support global fisheries production</article-title>
<source>Conserv Lett</source>
<year iso-8601-date="2019">2019</year>
<volume>12</volume>
<elocation-id>e12566</elocation-id>
<pub-id pub-id-type="doi">10.1111/conl.12566</pub-id></element-citation>
</ref>
<ref id="B22">
<label>22</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Duffy</surname>
<given-names>JE</given-names>
</name>
</person-group>
<article-title>Biodiversity and the functioning of seagrass ecosystems</article-title>
<source>Mar Ecol Prog Ser</source>
<year iso-8601-date="2006">2006</year>
<volume>311</volume>
<fpage>233</fpage>
<lpage>50</lpage>
<pub-id pub-id-type="doi">10.3354/meps311233</pub-id></element-citation>
</ref>
<ref id="B23">
<label>23</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Papenbrock</surname>
<given-names>J</given-names>
</name>
</person-group>
<article-title>Highlights in seagrasses’ phylogeny, physiology, and metabolism: What makes them special?</article-title>
<source>Int Sch Res Not</source>
<year iso-8601-date="2012">2012</year>
<volume>2012</volume>
<elocation-id>103892</elocation-id>
<pub-id pub-id-type="doi">10.5402/2012/103892</pub-id></element-citation>
</ref>
<ref id="B24">
<label>24</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Orth</surname>
<given-names>RJ</given-names>
</name>
<name>
<surname>Carruthers</surname>
<given-names>TJB</given-names>
</name>
<name>
<surname>Dennison</surname>
<given-names>W</given-names>
</name>
<name>
<surname>Duarte</surname>
<given-names>CM</given-names>
</name>
<name>
<surname>Fourqurean</surname>
<given-names>JW</given-names>
</name>
<name>
<surname>Heck</surname>
<given-names>KL</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>A global crisis for seagrass ecosystems</article-title>
<source>Bioscience</source>
<year iso-8601-date="2006">2006</year>
<volume>56</volume>
<fpage>987</fpage>
<lpage>96</lpage>
<pub-id pub-id-type="doi">10.1641/0006-3568(2006)56[987:AGCFSE]2.0.CO;2</pub-id></element-citation>
</ref>
<ref id="B25">
<label>25</label>
<element-citation publication-type="book">
<person-group person-group-type="author">
<name>
<surname>Jeyapragash</surname>
<given-names>D</given-names>
</name>
<name>
<surname>Saravanakumar</surname>
<given-names>A</given-names>
</name>
<name>
<surname>Yosuva</surname>
<given-names>M</given-names>
</name>
</person-group>
<article-title>Seagrass metabolomics: a new insight towards marine based drug discovery</article-title>
<person-group person-group-type="editor">
<name>
<surname>Zhan</surname>
<given-names>X</given-names>
</name>
</person-group>
<source>Metabolomics</source>
<publisher-loc>Rijeka</publisher-loc>
<publisher-name>IntechOpen</publisher-name>
<year iso-8601-date="2021">2021</year>
</element-citation>
</ref>
<ref id="B26">
<label>26</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Ugarelli</surname>
<given-names>K</given-names>
</name>
<name>
<surname>Chakrabarti</surname>
<given-names>S</given-names>
</name>
<name>
<surname>Laas</surname>
<given-names>P</given-names>
</name>
<name>
<surname>Stingl</surname>
<given-names>U</given-names>
</name>
</person-group>
<article-title>The seagrass holobiont and its microbiome</article-title>
<source>Microorganisms</source>
<year iso-8601-date="2017">2017</year>
<volume>5</volume>
<elocation-id>81</elocation-id>
<pub-id pub-id-type="doi">10.3390/microorganisms5040081</pub-id><pub-id pub-id-type="pmid">29244764</pub-id><pub-id pub-id-type="pmcid">PMC5748590</pub-id></element-citation>
</ref>
<ref id="B27">
<label>27</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Sandoval-Gil</surname>
<given-names>JM</given-names>
</name>
<name>
<surname>Ruiz</surname>
<given-names>JM</given-names>
</name>
<name>
<surname>Marín-Guirao</surname>
<given-names>L</given-names>
</name>
</person-group>
<article-title>Advances in understanding multilevel responses of seagrasses to hypersalinity</article-title>
<source>Mar Environ Res</source>
<year iso-8601-date="2023">2023</year>
<volume>183</volume>
<elocation-id>105809</elocation-id>
<pub-id pub-id-type="doi">10.1016/j.marenvres.2022.105809</pub-id><pub-id pub-id-type="pmid">36435174</pub-id></element-citation>
</ref>
<ref id="B28">
<label>28</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Short</surname>
<given-names>F</given-names>
</name>
<name>
<surname>Carruthers</surname>
<given-names>T</given-names>
</name>
<name>
<surname>Dennison</surname>
<given-names>W</given-names>
</name>
<name>
<surname>Waycott</surname>
<given-names>M</given-names>
</name>
</person-group>
<article-title>Global seagrass distribution and diversity: a bioregional model</article-title>
<source>J Exp Mar Bio Ecol</source>
<year iso-8601-date="2007">2007</year>
<volume>350</volume>
<fpage>3</fpage>
<lpage>20</lpage>
<pub-id pub-id-type="doi">10.1016/j.jembe.2007.06.012</pub-id></element-citation>
</ref>
<ref id="B29">
<label>29</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Ondiviela</surname>
<given-names>B</given-names>
</name>
<name>
<surname>Losada</surname>
<given-names>IJ</given-names>
</name>
<name>
<surname>Lara</surname>
<given-names>JL</given-names>
</name>
<name>
<surname>Maza</surname>
<given-names>M</given-names>
</name>
<name>
<surname>Galván</surname>
<given-names>C</given-names>
</name>
<name>
<surname>Bouma</surname>
<given-names>TJ</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>The role of seagrasses in coastal protection in a changing climate</article-title>
<source>Coast Eng</source>
<year iso-8601-date="2014">2014</year>
<volume>87</volume>
<fpage>158</fpage>
<lpage>68</lpage>
<pub-id pub-id-type="doi">10.1016/j.coastaleng.2013.11.005</pub-id></element-citation>
</ref>
<ref id="B30">
<label>30</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Rengasamy</surname>
<given-names>KRR</given-names>
</name>
<name>
<surname>Sadeer</surname>
<given-names>NB</given-names>
</name>
<name>
<surname>Zengin</surname>
<given-names>G</given-names>
</name>
<name>
<surname>Mahomoodally</surname>
<given-names>MF</given-names>
</name>
<name>
<surname>Cziáky</surname>
<given-names>Z</given-names>
</name>
<name>
<surname>Jekő</surname>
<given-names>J</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>Biopharmaceutical potential, chemical profile and <italic>in silico</italic> study of the seagrass– <italic>Syringodium isoetifolium</italic> (Asch.) Dandy</article-title>
<source>S Afr J Bot</source>
<year iso-8601-date="2019">2019</year>
<volume>127</volume>
<fpage>167</fpage>
<lpage>75</lpage>
<pub-id pub-id-type="doi">10.1016/j.sajb.2019.08.043</pub-id></element-citation>
</ref>
<ref id="B31">
<label>31</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>McKenzie</surname>
<given-names>LJ</given-names>
</name>
<name>
<surname>Nordlund</surname>
<given-names>LM</given-names>
</name>
<name>
<surname>Jones</surname>
<given-names>BL</given-names>
</name>
<name>
<surname>Cullen-Unsworth</surname>
<given-names>LC</given-names>
</name>
<name>
<surname>Roelfsema</surname>
<given-names>C</given-names>
</name>
<name>
<surname>Unsworth</surname>
<given-names>RKF</given-names>
</name>
</person-group>
<article-title>The global distribution of seagrass meadows</article-title>
<source>Environ Res Lett</source>
<year iso-8601-date="2020">2020</year>
<volume>15</volume>
<elocation-id>074041</elocation-id>
<pub-id pub-id-type="doi">10.1088/1748-9326/ab7d06</pub-id></element-citation>
</ref>
<ref id="B32">
<label>32</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Henrickson</surname>
<given-names>SE</given-names>
</name>
<name>
<surname>Wong</surname>
<given-names>T</given-names>
</name>
<name>
<surname>Allen</surname>
<given-names>P</given-names>
</name>
<name>
<surname>Ford</surname>
<given-names>T</given-names>
</name>
<name>
<surname>Epstein</surname>
<given-names>PR</given-names>
</name>
</person-group>
<article-title>Marine swimming-related illness: implications for monitoring and environmental policy</article-title>
<source>Environ Health Perspect</source>
<year iso-8601-date="2001">2001</year>
<volume>109</volume>
<fpage>645</fpage>
<lpage>50</lpage>
<pub-id pub-id-type="doi">10.1289/ehp.01109645</pub-id><pub-id pub-id-type="pmid">11485861</pub-id><pub-id pub-id-type="pmcid">PMC1240366</pub-id></element-citation>
</ref>
<ref id="B33">
<label>33</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Lamb</surname>
<given-names>JB</given-names>
</name>
<name>
<surname>van de Water</surname>
<given-names>JA</given-names>
</name>
<name>
<surname>Bourne</surname>
<given-names>DG</given-names>
</name>
<name>
<surname>Altier</surname>
<given-names>C</given-names>
</name>
<name>
<surname>Hein</surname>
<given-names>MY</given-names>
</name>
<name>
<surname>Fiorenza</surname>
<given-names>EA</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>Seagrass ecosystems reduce exposure to bacterial pathogens of humans, fishes, and invertebrates</article-title>
<source>Science</source>
<year iso-8601-date="2017">2017</year>
<volume>355</volume>
<fpage>731</fpage>
<lpage>3</lpage>
<pub-id pub-id-type="doi">10.1126/science.aal1956</pub-id><pub-id pub-id-type="pmid">28209895</pub-id></element-citation>
</ref>
<ref id="B34">
<label>34</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Ascioti</surname>
<given-names>FA</given-names>
</name>
<name>
<surname>Mangano</surname>
<given-names>MC</given-names>
</name>
<name>
<surname>Marcianò</surname>
<given-names>C</given-names>
</name>
<name>
<surname>Sarà</surname>
<given-names>G</given-names>
</name>
</person-group>
<article-title>The sanitation service of seagrasses – dependencies and implications for the estimation of avoided costs</article-title>
<source>Ecosyst Serv</source>
<year iso-8601-date="2022">2022</year>
<volume>54</volume>
<elocation-id>101418</elocation-id>
<pub-id pub-id-type="doi">10.1016/j.ecoser.2022.101418</pub-id></element-citation>
</ref>
<ref id="B35">
<label>35</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Zhao</surname>
<given-names>L</given-names>
</name>
<name>
<surname>Ru</surname>
<given-names>S</given-names>
</name>
<name>
<surname>He</surname>
<given-names>J</given-names>
</name>
<name>
<surname>Zhang</surname>
<given-names>Z</given-names>
</name>
<name>
<surname>Song</surname>
<given-names>X</given-names>
</name>
<name>
<surname>Wang</surname>
<given-names>D</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>Eelgrass (<italic>Zostera marina</italic>) and its epiphytic bacteria facilitate the sinking of microplastics in the seawater</article-title>
<source>Environ Pollut</source>
<year iso-8601-date="2022">2022</year>
<volume>292</volume>
<elocation-id>118337</elocation-id>
<pub-id pub-id-type="doi">10.1016/j.envpol.2021.118337</pub-id><pub-id pub-id-type="pmid">34644624</pub-id></element-citation>
</ref>
<ref id="B36">
<label>36</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Reusch</surname>
<given-names>TBH</given-names>
</name>
<name>
<surname>Schubert</surname>
<given-names>PR</given-names>
</name>
<name>
<surname>Marten</surname>
<given-names>SM</given-names>
</name>
<name>
<surname>Gill</surname>
<given-names>D</given-names>
</name>
<name>
<surname>Karez</surname>
<given-names>R</given-names>
</name>
<name>
<surname>Busch</surname>
<given-names>K</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>Lower <italic>Vibrio</italic> spp. abundances in <italic>Zostera marina</italic> leaf canopies suggest a novel ecosystem function for temperate seagrass beds</article-title>
<source>Mar Biol</source>
<year iso-8601-date="2021">2021</year>
<volume>168</volume>
<elocation-id>149</elocation-id>
<pub-id pub-id-type="doi">10.1007/s00227-021-03963-3</pub-id></element-citation>
</ref>
<ref id="B37">
<label>37</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Alsaffar</surname>
<given-names>Z</given-names>
</name>
<name>
<surname>Pearman</surname>
<given-names>JK</given-names>
</name>
<name>
<surname>Cúrdia</surname>
<given-names>J</given-names>
</name>
<name>
<surname>Ellis</surname>
<given-names>J</given-names>
</name>
<name>
<surname>Calleja</surname>
<given-names>ML</given-names>
</name>
<name>
<surname>Ruiz-Compean</surname>
<given-names>P</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>The role of seagrass vegetation and local environmental conditions in shaping benthic bacterial and macroinvertebrate communities in a tropical coastal lagoon</article-title>
<source>Sci Rep</source>
<year iso-8601-date="2020">2020</year>
<volume>10</volume>
<elocation-id>13550</elocation-id>
<pub-id pub-id-type="doi">10.1038/s41598-020-70318-1</pub-id><pub-id pub-id-type="pmid">32782295</pub-id><pub-id pub-id-type="pmcid">PMC7419567</pub-id></element-citation>
</ref>
<ref id="B38">
<label>38</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Tarquinio</surname>
<given-names>F</given-names>
</name>
<name>
<surname>Hyndes</surname>
<given-names>GA</given-names>
</name>
<name>
<surname>Laverock</surname>
<given-names>B</given-names>
</name>
<name>
<surname>Koenders</surname>
<given-names>A</given-names>
</name>
<name>
<surname>Säwström</surname>
<given-names>C</given-names>
</name>
</person-group>
<article-title>The seagrass holobiont: understanding seagrass-bacteria interactions and their role in seagrass ecosystem functioning</article-title>
<source>FEMS Microbiol Lett</source>
<year iso-8601-date="2019">2019</year>
<volume>366</volume>
<elocation-id>fnz057</elocation-id>
<pub-id pub-id-type="doi">10.1093/femsle/fnz057</pub-id><pub-id pub-id-type="pmid">30883643</pub-id></element-citation>
</ref>
<ref id="B39">
<label>39</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Conte</surname>
<given-names>C</given-names>
</name>
<name>
<surname>Rotini</surname>
<given-names>A</given-names>
</name>
<name>
<surname>Manfra</surname>
<given-names>L</given-names>
</name>
<name>
<surname>D’Andrea</surname>
<given-names>MM</given-names>
</name>
<name>
<surname>Winters</surname>
<given-names>G</given-names>
</name>
<name>
<surname>Migliore</surname>
<given-names>L</given-names>
</name>
</person-group>
<article-title>The seagrass holobiont: What we know and what we still need to disclose for its possible use as an ecological indicator</article-title>
<source>Water</source>
<year iso-8601-date="2021">2021</year>
<volume>13</volume>
<elocation-id>406</elocation-id>
<pub-id pub-id-type="doi">10.3390/w13040406</pub-id></element-citation>
</ref>
<ref id="B40">
<label>40</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Mohr</surname>
<given-names>W</given-names>
</name>
<name>
<surname>Lehnen</surname>
<given-names>N</given-names>
</name>
<name>
<surname>Ahmerkamp</surname>
<given-names>S</given-names>
</name>
<name>
<surname>Marchant</surname>
<given-names>HK</given-names>
</name>
<name>
<surname>Graf</surname>
<given-names>JS</given-names>
</name>
<name>
<surname>Tschitschko</surname>
<given-names>B</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>Terrestrial-type nitrogen-fixing symbiosis between seagrass and a marine bacterium</article-title>
<source>Nature</source>
<year iso-8601-date="2021">2021</year>
<volume>600</volume>
<fpage>105</fpage>
<lpage>9</lpage>
<pub-id pub-id-type="doi">10.1038/s41586-021-04063-4</pub-id><pub-id pub-id-type="pmid">34732889</pub-id><pub-id pub-id-type="pmcid">PMC8636270</pub-id></element-citation>
</ref>
<ref id="B41">
<label>41</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Iqbal</surname>
<given-names>MM</given-names>
</name>
<name>
<surname>Nishimura</surname>
<given-names>M</given-names>
</name>
<name>
<surname>Haider</surname>
<given-names>MN</given-names>
</name>
<name>
<surname>Sano</surname>
<given-names>M</given-names>
</name>
<name>
<surname>Ijichi</surname>
<given-names>M</given-names>
</name>
<name>
<surname>Kogure</surname>
<given-names>K</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>Diversity and composition of microbial communities in an eelgrass (<italic>Zostera marina</italic>) bed in Tokyo Bay, Japan</article-title>
<source>Microbes Environ</source>
<year iso-8601-date="2021">2021</year>
<volume>36</volume>
<elocation-id>ME21037</elocation-id>
<pub-id pub-id-type="doi">10.1264/jsme2.ME21037</pub-id><pub-id pub-id-type="pmid">34645731</pub-id><pub-id pub-id-type="pmcid">PMC8674447</pub-id></element-citation>
</ref>
<ref id="B42">
<label>42</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Trevathan-Tackett</surname>
<given-names>SM</given-names>
</name>
<name>
<surname>Lane</surname>
<given-names>AL</given-names>
</name>
<name>
<surname>Bishop</surname>
<given-names>N</given-names>
</name>
<name>
<surname>Ross</surname>
<given-names>C</given-names>
</name>
</person-group>
<article-title>Metabolites derived from the tropical seagrass <italic>Thalassia testudinum</italic> are bioactive against pathogenic <italic>Labyrinthula</italic> sp</article-title>
<source>Aquat Bot</source>
<year iso-8601-date="2015">2015</year>
<volume>122</volume>
<fpage>1</fpage>
<lpage>8</lpage>
<pub-id pub-id-type="doi">10.1016/j.aquabot.2014.12.005</pub-id></element-citation>
</ref>
<ref id="B43">
<label>43</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Tarquinio</surname>
<given-names>F</given-names>
</name>
<name>
<surname>Attlan</surname>
<given-names>O</given-names>
</name>
<name>
<surname>Vanderklift</surname>
<given-names>MA</given-names>
</name>
<name>
<surname>Berry</surname>
<given-names>O</given-names>
</name>
<name>
<surname>Bissett</surname>
<given-names>A</given-names>
</name>
</person-group>
<article-title>Distinct endophytic bacterial communities inhabiting seagrass seeds</article-title>
<source>Front Microbiol</source>
<year iso-8601-date="2021">2021</year>
<volume>12</volume>
<elocation-id>703014</elocation-id>
<pub-id pub-id-type="doi">10.3389/fmicb.2021.703014</pub-id><pub-id pub-id-type="pmid">34621247</pub-id><pub-id pub-id-type="pmcid">PMC8491609</pub-id></element-citation>
</ref>
<ref id="B44">
<label>44</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Hurtado-McCormick</surname>
<given-names>V</given-names>
</name>
<name>
<surname>Kahlke</surname>
<given-names>T</given-names>
</name>
<name>
<surname>Petrou</surname>
<given-names>K</given-names>
</name>
<name>
<surname>Jeffries</surname>
<given-names>T</given-names>
</name>
<name>
<surname>Ralph</surname>
<given-names>PJ</given-names>
</name>
<name>
<surname>Seymour</surname>
<given-names>JR</given-names>
</name>
</person-group>
<article-title>Regional and microenvironmental scale characterization of the <italic>Zostera muelleri</italic> seagrass Microbiome</article-title>
<source>Front Microbiol</source>
<year iso-8601-date="2019">2019</year>
<volume>10</volume>
<elocation-id>1011</elocation-id>
<comment>Erratum in: Front Microbiol. 2021;12:642964.</comment>
<pub-id pub-id-type="doi">10.3389/fmicb.2019.01011</pub-id><pub-id pub-id-type="pmid">31139163</pub-id><pub-id pub-id-type="pmcid">PMC6527750</pub-id></element-citation>
</ref>
<ref id="B45">
<label>45</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Conte</surname>
<given-names>C</given-names>
</name>
<name>
<surname>Apostolaki</surname>
<given-names>ET</given-names>
</name>
<name>
<surname>Vizzini</surname>
<given-names>S</given-names>
</name>
<name>
<surname>Migliore</surname>
<given-names>L</given-names>
</name>
</person-group>
<article-title>A tight interaction between the native seagrass <italic>Cymodocea nodosa</italic> and the exotic <italic>Halophila stipulacea</italic> in the Aegean Sea highlights seagrass holobiont variations</article-title>
<source>Plants (Basel)</source>
<year iso-8601-date="2023">2023</year>
<volume>12</volume>
<elocation-id>350</elocation-id>
<pub-id pub-id-type="doi">10.3390/plants12020350</pub-id><pub-id pub-id-type="pmid">36679063</pub-id><pub-id pub-id-type="pmcid">PMC9863530</pub-id></element-citation>
</ref>
<ref id="B46">
<label>46</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Mohapatra</surname>
<given-names>M</given-names>
</name>
<name>
<surname>Manu</surname>
<given-names>S</given-names>
</name>
<name>
<surname>Dash</surname>
<given-names>SP</given-names>
</name>
<name>
<surname>Rastogi</surname>
<given-names>G</given-names>
</name>
</person-group>
<article-title>Seagrasses and local environment control the bacterial community structure and carbon substrate utilization in brackish sediments</article-title>
<source>J Environ Manage</source>
<year iso-8601-date="2022">2022</year>
<volume>314</volume>
<elocation-id>115013</elocation-id>
<pub-id pub-id-type="doi">10.1016/j.jenvman.2022.115013</pub-id><pub-id pub-id-type="pmid">35447445</pub-id></element-citation>
</ref>
<ref id="B47">
<label>47</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Banister</surname>
<given-names>RB</given-names>
</name>
<name>
<surname>Schwarz</surname>
<given-names>MT</given-names>
</name>
<name>
<surname>Fine</surname>
<given-names>M</given-names>
</name>
<name>
<surname>Ritchie</surname>
<given-names>KB</given-names>
</name>
<name>
<surname>Muller</surname>
<given-names>EM</given-names>
</name>
</person-group>
<article-title>Instability and stasis among the microbiome of seagrass leaves, roots and rhizomes, and nearby sediments within a natural pH gradient</article-title>
<source>Microb Ecol</source>
<year iso-8601-date="2022">2022</year>
<volume>84</volume>
<fpage>703</fpage>
<lpage>16</lpage>
<comment>Erratum in: Microb Ecol. 2023;85:1634.</comment>
<pub-id pub-id-type="doi">10.1007/s00248-021-01867-9</pub-id><pub-id pub-id-type="pmid">34596709</pub-id><pub-id pub-id-type="pmcid">PMC9622545</pub-id></element-citation>
</ref>
<ref id="B48">
<label>48</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Vasarri</surname>
<given-names>M</given-names>
</name>
<name>
<surname>De</surname>
<given-names>Biasi AM</given-names>
</name>
<name>
<surname>Barletta</surname>
<given-names>E</given-names>
</name>
<name>
<surname>Pretti</surname>
<given-names>C</given-names>
</name>
<name>
<surname>Degl’Innocenti</surname>
<given-names>D</given-names>
</name>
</person-group>
<article-title>An overview of new insights into the benefits of the seagrass <italic>Posidonia oceanica</italic> for human health</article-title>
<source>Mar Drugs</source>
<year iso-8601-date="2021">2021</year>
<volume>19</volume>
<elocation-id>476</elocation-id>
<pub-id pub-id-type="doi">10.3390/md19090476</pub-id><pub-id pub-id-type="pmid">34564138</pub-id><pub-id pub-id-type="pmcid">PMC8470915</pub-id></element-citation>
</ref>
<ref id="B49">
<label>49</label>
<element-citation publication-type="book">
<person-group person-group-type="author">
<name>
<surname>Terrados</surname>
<given-names>J</given-names>
</name>
<name>
<surname>Borum</surname>
<given-names>J</given-names>
</name>
</person-group>
<article-title>Why are seagrasses important? - Goods and services provided by seagrass meadows</article-title>
<person-group person-group-type="editor">
<name>
<surname>Borum</surname>
<given-names>J</given-names>
</name>
<name>
<surname>Duarte</surname>
<given-names>CM</given-names>
</name>
<name>
<surname>Krause-Jensen</surname>
<given-names>D</given-names>
</name>
<name>
<surname>Greve</surname>
<given-names>TM</given-names>
</name>
</person-group>
<source>European seagrasses: an introduction to monitoring and management</source>
<comment>The M&amp;MS project; 2004. pp. 8–10.</comment>
</element-citation>
</ref>
<ref id="B50">
<label>50</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Zidorn</surname>
<given-names>C</given-names>
</name>
</person-group>
<article-title>Secondary metabolites of seagrasses (Alismatales and Potamogetonales; Alismatidae): chemical diversity, bioactivity, and ecological function</article-title>
<source>Phytochemistry</source>
<year iso-8601-date="2016">2016</year>
<volume>124</volume>
<fpage>5</fpage>
<lpage>28</lpage>
<pub-id pub-id-type="doi">10.1016/j.phytochem.2016.02.004</pub-id><pub-id pub-id-type="pmid">26880288</pub-id></element-citation>
</ref>
<ref id="B51">
<label>51</label>
<element-citation publication-type="book">
<person-group person-group-type="author">
<name>
<surname>Jiang</surname>
<given-names>Y</given-names>
</name>
<name>
<surname>Li</surname>
<given-names>Q</given-names>
</name>
<name>
<surname>Chen</surname>
<given-names>X</given-names>
</name>
<name>
<surname>Jiang</surname>
<given-names>C</given-names>
</name>
</person-group>
<article-title>Isolation and cultivation methods of actinobacteria</article-title>
<person-group person-group-type="editor">
<name>
<surname>Dhanasekaran</surname>
<given-names>D</given-names>
</name>
<name>
<surname>Jiang</surname>
<given-names>Y</given-names>
</name>
</person-group>
<source>Actinobacteria</source>
<publisher-loc>Rijeka</publisher-loc>
<publisher-name>IntechOpen</publisher-name>
<year iso-8601-date="2016">2016</year>
<pub-id pub-id-type="doi">10.5772/61457</pub-id></element-citation>
</ref>
<ref id="B52">
<label>52</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Wu</surname>
<given-names>H</given-names>
</name>
<name>
<surname>Chen</surname>
<given-names>W</given-names>
</name>
<name>
<surname>Wang</surname>
<given-names>G</given-names>
</name>
<name>
<surname>Dai</surname>
<given-names>S</given-names>
</name>
<name>
<surname>Zhou</surname>
<given-names>D</given-names>
</name>
<name>
<surname>Zhao</surname>
<given-names>H</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>Culture-dependent diversity of <italic>actinobacteria</italic> associated with seagrass (<italic>Thalassia hemprichii</italic>)</article-title>
<source>Afr J Microbiol Res</source>
<year iso-8601-date="2012">2012</year>
<volume>6</volume>
<fpage>87</fpage>
<lpage>94</lpage>
<pub-id pub-id-type="doi">10.5897/AJMR11.981</pub-id></element-citation>
</ref>
<ref id="B53">
<label>53</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Ravikumar</surname>
<given-names>S</given-names>
</name>
<name>
<surname>Gnanadesigan</surname>
<given-names>M</given-names>
</name>
<name>
<surname>Saravanan</surname>
<given-names>A</given-names>
</name>
<name>
<surname>Monisha</surname>
<given-names>N</given-names>
</name>
<name>
<surname>Brindha</surname>
<given-names>V</given-names>
</name>
<name>
<surname>Muthumari</surname>
<given-names>S</given-names>
</name>
</person-group>
<article-title>Antagonistic properties of seagrass associated <italic>Streptomyces</italic> sp. RAUACT-1: a source for anthraquinone rich compound</article-title>
<source>Asian Pac J Trop Med</source>
<year iso-8601-date="2012">2012</year>
<volume>5</volume>
<fpage>887</fpage>
<lpage>90</lpage>
<pub-id pub-id-type="doi">10.1016/S1995-7645(12)60165-5</pub-id><pub-id pub-id-type="pmid">23146803</pub-id></element-citation>
</ref>
<ref id="B54">
<label>54</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Jose</surname>
<given-names>PA</given-names>
</name>
<name>
<surname>sundari</surname>
<given-names>IS</given-names>
</name>
<name>
<surname>Sivakala</surname>
<given-names>KK</given-names>
</name>
<name>
<surname>Jebakumar</surname>
<given-names>SRD</given-names>
</name>
</person-group>
<article-title>Molecular phylogeny and plant growth promoting traits of endophytic bacteria isolated from roots of seagrass <italic>Cymodocea serrulata</italic></article-title>
<source>Indian J Geo-Mar Sci</source>
<year iso-8601-date="2014">2014</year>
<volume>43</volume>
<fpage>571</fpage>
<lpage>9</lpage>
</element-citation>
</ref>
<ref id="B55">
<label>55</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Bibi</surname>
<given-names>F</given-names>
</name>
<name>
<surname>Naseer</surname>
<given-names>MI</given-names>
</name>
<name>
<surname>Hassan</surname>
<given-names>AM</given-names>
</name>
<name>
<surname>Yasir</surname>
<given-names>M</given-names>
</name>
<name>
<surname>Al-Ghamdi</surname>
<given-names>AAK</given-names>
</name>
<name>
<surname>Azhar</surname>
<given-names>EI</given-names>
</name>
</person-group>
<article-title>Diversity and antagonistic potential of bacteria isolated from marine grass <italic>Halodule uninervis</italic></article-title>
<source>3 Biotech</source>
<year iso-8601-date="2018">2018</year>
<volume>8</volume>
<elocation-id>48</elocation-id>
<pub-id pub-id-type="doi">10.1007/s13205-017-1066-1</pub-id><pub-id pub-id-type="pmid">29354359</pub-id><pub-id pub-id-type="pmcid">PMC5752661</pub-id></element-citation>
</ref>
<ref id="B56">
<label>56</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Ettinger</surname>
<given-names>CL</given-names>
</name>
<name>
<surname>Eisen</surname>
<given-names>JA</given-names>
</name>
</person-group>
<article-title>Fungi, bacteria and oomycota opportunistically isolated from the seagrass, <italic>Zostera marina</italic></article-title>
<source>PLoS One</source>
<year iso-8601-date="2020">2020</year>
<volume>15</volume>
<elocation-id>e0236135</elocation-id>
<comment>Erratum in: PLoS One. 2021;16:e0251536.</comment>
<pub-id pub-id-type="doi">10.1371/journal.pone.0236135</pub-id><pub-id pub-id-type="pmid">32697800</pub-id><pub-id pub-id-type="pmcid">PMC7375540</pub-id></element-citation>
</ref>
<ref id="B57">
<label>57</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Cristianawati</surname>
<given-names>O</given-names>
</name>
<name>
<surname>Sibero</surname>
<given-names>MT</given-names>
</name>
<name>
<surname>Ayuningrum</surname>
<given-names>D</given-names>
</name>
<name>
<surname>Nuryadi</surname>
<given-names>H</given-names>
</name>
<name>
<surname>Syafitri</surname>
<given-names>E</given-names>
</name>
<name>
<surname>Riniarsih</surname>
<given-names>I</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>Screening of antibacterial activity of seagrass-associated bacteria from the North Java Sea, Indonesia against multidrug-resistant bacteria</article-title>
<source>Aquac Aquarium Conserv Legis</source>
<year iso-8601-date="2019">2019</year>
<volume>12</volume>
<fpage>1054</fpage>
<lpage>64</lpage>
</element-citation>
</ref>
<ref id="B58">
<label>58</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Hamdy</surname>
<given-names>AHA</given-names>
</name>
<name>
<surname>El-Fiky</surname>
<given-names>NM</given-names>
</name>
<name>
<surname>El-Beih</surname>
<given-names>AA</given-names>
</name>
<name>
<surname>Mohammed</surname>
<given-names>MM</given-names>
</name>
<name>
<surname>Mettwally</surname>
<given-names>WS</given-names>
</name>
</person-group>
<article-title>Egyptian red sea seagrass as a source of biologically active secondary metabolites</article-title>
<source>Egypt Pharm J</source>
<year iso-8601-date="2020">2020</year>
<volume>19</volume>
<fpage>224</fpage>
<lpage>37</lpage>
<pub-id pub-id-type="doi">10.4103/epj.epj_57_19</pub-id></element-citation>
</ref>
<ref id="B59">
<label>59</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Ragupathi</surname>
<given-names>Raja Kannan R</given-names>
</name>
<name>
<surname>Arumugam</surname>
<given-names>R</given-names>
</name>
<name>
<surname>Anantharaman</surname>
<given-names>P</given-names>
</name>
</person-group>
<article-title>Chemical composition and antibacterial activity of Indian seagrasses against urinary tract pathogens</article-title>
<source>Food Chem</source>
<year iso-8601-date="2012">2012</year>
<volume>135</volume>
<fpage>2470</fpage>
<lpage>3</lpage>
<pub-id pub-id-type="doi">10.1016/j.foodchem.2012.07.070</pub-id><pub-id pub-id-type="pmid">22980830</pub-id></element-citation>
</ref>
<ref id="B60">
<label>60</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Engel</surname>
<given-names>S</given-names>
</name>
<name>
<surname>Puglisi</surname>
<given-names>MP</given-names>
</name>
<name>
<surname>Jensen</surname>
<given-names>PR</given-names>
</name>
<name>
<surname>Fenical</surname>
<given-names>W</given-names>
</name>
</person-group>
<article-title>Antimicrobial activities of extracts from tropical Atlantic marine plants against marine pathogens and saprophytes</article-title>
<source>Mar Biol</source>
<year iso-8601-date="2006">2006</year>
<volume>149</volume>
<fpage>991</fpage>
<lpage>1002</lpage>
<pub-id pub-id-type="doi">10.1007/s00227-006-0264-x</pub-id></element-citation>
</ref>
<ref id="B61">
<label>61</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Kim</surname>
<given-names>DH</given-names>
</name>
<name>
<surname>Mahomoodally</surname>
<given-names>MF</given-names>
</name>
<name>
<surname>Sadeer</surname>
<given-names>NB</given-names>
</name>
<name>
<surname>Seok</surname>
<given-names>PG</given-names>
</name>
<name>
<surname>Zengin</surname>
<given-names>G</given-names>
</name>
<name>
<surname>Palaniveloo</surname>
<given-names>K</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>Nutritional and bioactive potential of seagrasses: a review</article-title>
<source>S Afr J Bot</source>
<year iso-8601-date="2021">2021</year>
<volume>137</volume>
<fpage>216</fpage>
<lpage>27</lpage>
<pub-id pub-id-type="doi">10.1016/j.sajb.2020.10.018</pub-id></element-citation>
</ref>
<ref id="B62">
<label>62</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Gono</surname>
<given-names>CMP</given-names>
</name>
<name>
<surname>Ahmadi</surname>
<given-names>P</given-names>
</name>
<name>
<surname>Hertiani</surname>
<given-names>T</given-names>
</name>
<name>
<surname>Septiana</surname>
<given-names>E</given-names>
</name>
<name>
<surname>Putra</surname>
<given-names>MY</given-names>
</name>
<name>
<surname>Chianese</surname>
<given-names>G</given-names>
</name>
</person-group>
<article-title>A comprehensive update on the bioactive compounds from seagrasses</article-title>
<source>Mar Drugs</source>
<year iso-8601-date="2022">2022</year>
<volume>20</volume>
<elocation-id>406</elocation-id>
<pub-id pub-id-type="doi">10.3390/md20070406</pub-id><pub-id pub-id-type="pmid">35877699</pub-id><pub-id pub-id-type="pmcid">PMC9324380</pub-id></element-citation>
</ref>
<ref id="B63">
<label>63</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Jafriati</surname>
<given-names>J</given-names>
</name>
<name>
<surname>Hatta</surname>
<given-names>M</given-names>
</name>
<name>
<surname>Yuniar</surname>
<given-names>N</given-names>
</name>
<name>
<surname>Ade</surname>
<given-names>RJ</given-names>
</name>
<name>
<surname>Dwiyanti</surname>
<given-names>R</given-names>
</name>
<name>
<surname>Sabir</surname>
<given-names>M</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>
<italic>Thalassia hemprichii</italic> seagrass extract as antimicrobial and antioxidant potential on human: a mini review of the benefits of seagrass</article-title>
<source>J Biol Sci</source>
<year iso-8601-date="2019">2019</year>
<volume>19</volume>
<fpage>363</fpage>
<lpage>71</lpage>
<pub-id pub-id-type="doi">10.3923/jbs.2019.363.371</pub-id></element-citation>
</ref>
<ref id="B64">
<label>64</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Nur</surname>
<given-names>RM</given-names>
</name>
<name>
<surname>Nurafni</surname>
</name>
<name>
<surname>Koroy</surname>
<given-names>K</given-names>
</name>
<name>
<surname>Alwi</surname>
<given-names>D</given-names>
</name>
<name>
<surname>Wahab</surname>
<given-names>I</given-names>
</name>
<name>
<surname>Sulistiawati</surname>
<given-names>S</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>The antibacterial activity of seagrass <italic>Enhalus acoroides</italic> against <italic>Staphylococcus aureus</italic></article-title>
<source>IOP Conf Ser: Earth Environ Sci</source>
<year iso-8601-date="2021">2021</year>
<volume>890</volume>
<elocation-id>012013</elocation-id>
<pub-id pub-id-type="doi">10.1088/1755-1315/890/1/012013</pub-id></element-citation>
</ref>
<ref id="B65">
<label>65</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>De</surname>
<given-names>Vincenti L</given-names>
</name>
<name>
<surname>Glasenapp</surname>
<given-names>Y</given-names>
</name>
<name>
<surname>Cattò</surname>
<given-names>C</given-names>
</name>
<name>
<surname>Villa</surname>
<given-names>F</given-names>
</name>
<name>
<surname>Cappitelli</surname>
<given-names>F</given-names>
</name>
<name>
<surname>Papenbrock</surname>
<given-names>J</given-names>
</name>
</person-group>
<article-title>Hindering the formation and promoting the dispersion of medical biofilms: non-lethal effects of seagrass extracts</article-title>
<source>BMC Complement Altern Med</source>
<year iso-8601-date="2018">2018</year>
<volume>18</volume>
<elocation-id>168</elocation-id>
<pub-id pub-id-type="doi">10.1186/s12906-018-2232-7</pub-id><pub-id pub-id-type="pmid">29843708</pub-id><pub-id pub-id-type="pmcid">PMC5975390</pub-id></element-citation>
</ref>
<ref id="B66">
<label>66</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Sun</surname>
<given-names>Y</given-names>
</name>
<name>
<surname>Song</surname>
<given-names>Z</given-names>
</name>
<name>
<surname>Zhang</surname>
<given-names>H</given-names>
</name>
<name>
<surname>Liu</surname>
<given-names>P</given-names>
</name>
<name>
<surname>Hu</surname>
<given-names>X</given-names>
</name>
</person-group>
<article-title>Seagrass vegetation affect the vertical organization of microbial communities in sediment</article-title>
<source>Mar Environ Res</source>
<year iso-8601-date="2020">2020</year>
<volume>162</volume>
<elocation-id>105174</elocation-id>
<pub-id pub-id-type="doi">10.1016/j.marenvres.2020.105174</pub-id><pub-id pub-id-type="pmid">33099080</pub-id></element-citation>
</ref>
<ref id="B67">
<label>67</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Crump</surname>
<given-names>BC</given-names>
</name>
<name>
<surname>Wojahn</surname>
<given-names>JM</given-names>
</name>
<name>
<surname>Tomas</surname>
<given-names>F</given-names>
</name>
<name>
<surname>Mueller</surname>
<given-names>RS</given-names>
</name>
</person-group>
<article-title>Metatranscriptomics and amplicon sequencing reveal mutualisms in seagrass microbiomes</article-title>
<source>Front Microbiol</source>
<year iso-8601-date="2018">2018</year>
<volume>9</volume>
<elocation-id>388</elocation-id>
<pub-id pub-id-type="doi">10.3389/fmicb.2018.00388</pub-id><pub-id pub-id-type="pmid">29599758</pub-id><pub-id pub-id-type="pmcid">PMC5863793</pub-id></element-citation>
</ref>
<ref id="B68">
<label>68</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Rotini</surname>
<given-names>A</given-names>
</name>
<name>
<surname>Conte</surname>
<given-names>C</given-names>
</name>
<name>
<surname>Seveso</surname>
<given-names>D</given-names>
</name>
<name>
<surname>Montano</surname>
<given-names>S</given-names>
</name>
<name>
<surname>Galli</surname>
<given-names>P</given-names>
</name>
<name>
<surname>Vai</surname>
<given-names>M</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>Daily variation of the associated microbial community and the Hsp60 expression in the Maldivian seagrass <italic>Thalassia hemprichii</italic></article-title>
<source>J Sea Res</source>
<year iso-8601-date="2020">2020</year>
<volume>156</volume>
<elocation-id>101835</elocation-id>
<pub-id pub-id-type="doi">10.1016/j.seares.2019.101835</pub-id></element-citation>
</ref>
<ref id="B69">
<label>69</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Ugarelli</surname>
<given-names>K</given-names>
</name>
<name>
<surname>Laas</surname>
<given-names>P</given-names>
</name>
<name>
<surname>Stingl</surname>
<given-names>U</given-names>
</name>
</person-group>
<article-title>The microbial communities of leaves and roots associated with turtle grass (<italic>Thalassia testudinum</italic>) and manatee grass (<italic>Syringodium filliforme</italic>) are distinct from seawater and sediment communities, but are similar between species and sampling sites</article-title>
<source>Microorganisms</source>
<year iso-8601-date="2018">2018</year>
<volume>7</volume>
<elocation-id>4</elocation-id>
<pub-id pub-id-type="doi">10.3390/microorganisms7010004</pub-id><pub-id pub-id-type="pmid">30587804</pub-id><pub-id pub-id-type="pmcid">PMC6352278</pub-id></element-citation>
</ref>
<ref id="B70">
<label>70</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Jensen</surname>
<given-names>SI</given-names>
</name>
<name>
<surname>Kühl</surname>
<given-names>M</given-names>
</name>
<name>
<surname>Priemé</surname>
<given-names>A</given-names>
</name>
</person-group>
<article-title>Different bacterial communities associated with the roots and bulk sediment of the seagrass <italic>Zostera marina</italic></article-title>
<source>FEMS Microbiol Ecol</source>
<year iso-8601-date="2007">2007</year>
<volume>62</volume>
<fpage>108</fpage>
<lpage>17</lpage>
<pub-id pub-id-type="doi">10.1111/j.1574-6941.2007.00373.x</pub-id><pub-id pub-id-type="pmid">17825072</pub-id></element-citation>
</ref>
<ref id="B71">
<label>71</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Damayanti</surname>
<given-names>V</given-names>
</name>
<name>
<surname>Rachma</surname>
<given-names>RN</given-names>
</name>
<name>
<surname>Santoso</surname>
<given-names>I</given-names>
</name>
<name>
<surname>Yasman</surname>
<given-names>Y</given-names>
</name>
<name>
<surname>Maryanto</surname>
<given-names>AE</given-names>
</name>
</person-group>
<article-title>Fermentation of antimicrobial substances of <italic>Streptomyces</italic> sp. BCy isolated from seagrass <italic>Cymodocearotundata</italic> using two different media</article-title>
<source>AIP Conf Proc</source>
<year iso-8601-date="2018">2018</year>
<volume>2023</volume>
<elocation-id>020143</elocation-id>
<pub-id pub-id-type="doi">10.1063/1.5064140</pub-id></element-citation>
</ref>
<ref id="B72">
<label>72</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Almaary</surname>
<given-names>KS</given-names>
</name>
<name>
<surname>Alharbi</surname>
<given-names>NS</given-names>
</name>
<name>
<surname>Kadaikunnan</surname>
<given-names>S</given-names>
</name>
<name>
<surname>Khaled</surname>
<given-names>JM</given-names>
</name>
<name>
<surname>Rajivgandhi</surname>
<given-names>G</given-names>
</name>
<name>
<surname>Ramachandran</surname>
<given-names>G</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>Anti-bacterial effect of marine sea grasses mediated endophytic actinomycetes against <italic>K. pneumoniae</italic></article-title>
<source>J King Saud Univ Sci</source>
<year iso-8601-date="2021">2021</year>
<volume>33</volume>
<elocation-id>101528</elocation-id>
<pub-id pub-id-type="doi">10.1016/j.jksus.2021.101528</pub-id></element-citation>
</ref>
<ref id="B73">
<label>73</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Inaba</surname>
<given-names>N</given-names>
</name>
<name>
<surname>Trainer</surname>
<given-names>VL</given-names>
</name>
<name>
<surname>Onishi</surname>
<given-names>Y</given-names>
</name>
<name>
<surname>Ishii</surname>
<given-names>KI</given-names>
</name>
<name>
<surname>Wyllie-Echeverria</surname>
<given-names>S</given-names>
</name>
<name>
<surname>Imai</surname>
<given-names>I</given-names>
</name>
</person-group>
<article-title>Algicidal and growth-inhibiting bacteria associated with seagrass and macroalgae beds in Puget Sound, WA, USA</article-title>
<source>Harmful Algae</source>
<year iso-8601-date="2017">2017</year>
<volume>62</volume>
<fpage>136</fpage>
<lpage>47</lpage>
<pub-id pub-id-type="doi">10.1016/j.hal.2016.04.004</pub-id><pub-id pub-id-type="pmid">28118888</pub-id></element-citation>
</ref>
<ref id="B74">
<label>74</label>
<element-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Martin</surname>
<given-names>BC</given-names>
</name>
<name>
<surname>Gleeson</surname>
<given-names>D</given-names>
</name>
<name>
<surname>Statton</surname>
<given-names>J</given-names>
</name>
<name>
<surname>Siebers</surname>
<given-names>AR</given-names>
</name>
<name>
<surname>Grierson</surname>
<given-names>P</given-names>
</name>
<name>
<surname>Ryan</surname>
<given-names>MH</given-names>
</name>
<etal>et al.</etal>
</person-group>
<article-title>Low light availability alters root exudation and reduces putative beneficial microorganisms in seagrass roots</article-title>
<source>Front Microbiol</source>
<year iso-8601-date="2018">2018</year>
<volume>8</volume>
<elocation-id>2667</elocation-id>
<pub-id pub-id-type="doi">10.3389/fmicb.2017.02667</pub-id><pub-id pub-id-type="pmid">29375529</pub-id><pub-id pub-id-type="pmcid">PMC5768916</pub-id></element-citation>
</ref>
</ref-list>
</back>
</article>