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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Immunol.</journal-id>
<journal-title>Frontiers in Immunology</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Immunol.</abbrev-journal-title>
<issn pub-type="epub">1664-3224</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fimmu.2021.762082</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Immunology</subject>
<subj-group>
<subject>Editorial</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Editorial: Aquatic Invertebrate Immunity Against Infectious Diseases</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Perazzolo</surname>
<given-names>Luciane Maria</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/928749"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Li</surname>
<given-names>Chaozheng</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/571941/overview"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Somboonwiwat</surname>
<given-names>Kunlaya</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/421452"/>
</contrib>
</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>Laboratory of Immunology Applied to Aquaculture, Department of Cell Biology, Embryology and Genetics, Federal University of Santa Catarina</institution>, <addr-line>Florian&#xf3;polis</addr-line>, <country>Brazil</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai)/ State Key Laboratory of Biocontrol, School of Marine Sciences, Sun Yat-Sen University</institution>, <addr-line>Guangzhou</addr-line>, <country>China</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>Center of Excellence for Molecular Biology and Genomics of Shrimp, Department of Biochemistry, Faculty of Science, Chulalongkorn University</institution>, <addr-line>Bangkok</addr-line>, <country>Thailand</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited and reviewed by: Miki Nakao, Kyushu University, Japan</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Luciane Maria Perazzolo, <email xlink:href="mailto:l.m.perazzolo@ufsc.br">l.m.perazzolo@ufsc.br</email>
</p>
</fn>
<fn fn-type="other" id="fn002">
<p>This article was submitted to Comparative Immunology, a section of the journal Frontiers in Immunology</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>22</day>
<month>09</month>
<year>2021</year>
</pub-date>
<pub-date pub-type="collection">
<year>2021</year>
</pub-date>
<volume>12</volume>
<elocation-id>762082</elocation-id>
<history>
<date date-type="received">
<day>20</day>
<month>08</month>
<year>2021</year>
</date>
<date date-type="accepted">
<day>06</day>
<month>09</month>
<year>2021</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2021 Perazzolo, Li and Somboonwiwat</copyright-statement>
<copyright-year>2021</copyright-year>
<copyright-holder>Perazzolo, Li and Somboonwiwat</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<related-article id="RA1" related-article-type="commentary-article" xlink:href="https://www.frontiersin.org/research-topics/13598/aquatic-invertebrate-immunity-against-infectious-diseases" ext-link-type="uri">Editorial on the Research Topic <article-title>Aquatic Invertebrate Immunity Against Infectious Diseases</article-title>
</related-article>
<kwd-group>
<kwd>farmed aquatic invertebrates</kwd>
<kwd>host-pathogen-microbiota interactions</kwd>
<kwd>antiviral immunity</kwd>
<kwd>antimicrobial defense</kwd>
<kwd>crustaceans</kwd>
<kwd>mollusks</kwd>
</kwd-group>
<counts>
<fig-count count="0"/>
<table-count count="0"/>
<equation-count count="0"/>
<ref-count count="5"/>
<page-count count="3"/>
<word-count count="1299"/>
</counts>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<title> </title>
<p>Aquatic invertebrates farming represents an important trade in the food supply of the global economy. Despite the expanding commercial relevance of shellfish farming, emerging diseases have caused catastrophic losses by limiting aquaculture sustainability. Crustaceans and mollusks cultures face numerous challenges, notably viral, bacterial, and fungal infections. Invertebrates lack classical adaptive immunity and only rely on innate responses to protect them from pathogens. Shrimp, oysters, scallops, mussels, crayfish, and crabs are among the most valuable farmed invertebrates. Understanding interactions between its immunity and pathogens is critical to controlling farming diseases. In light of this, nowadays, research on shellfish immunity has received remarkable efforts to unravel the molecular mechanisms that drive the immune responses against pathogens and, ultimately, to reach new non-antibody-based therapies to protect both cultivated and wild species. This Research Topic compiles 13 articles, including original studies and reviews by renowned scientists and research groups, which integrate current progress and understanding regarding antiviral and antimicrobial defenses triggered by aquatic invertebrates to control infectious diseases.</p>
</sec>
<sec id="s2">
<title>Pathogen Recognition and Antimicrobial Defenses</title>
<p>Like other invertebrates, shellfish possess cellular and humoral defenses to protect them against pathogens. However, defense responses must be tightly regulated to allow the destruction of pathogens without harming the host. In this Research Topic, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fimmu.2021.667787">Bouallegui</ext-link> reviews crayfish immunity, addressing plague disease, molecular aspects of hematopoiesis, and highlighting cell-mediated responses associated with Extracellular trap cell death (ETosis). During pathogen recognition, the host&#x2019;s pattern recognition receptors/proteins (PRRs/PRPs) recognize and bind to microbial/pathogen-associated molecular patterns (MAMPs/PAMPs), inducing PAMP-triggered immunity (PTI). C-type lectins are crucial PRPs for immobilizing microorganisms by agglutination; shrimp lectins are also implicated in antibacterial and antiviral defenses (<xref ref-type="bibr" rid="B1">1</xref>). <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fimmu.2021.661823">Qiu et&#xa0;al.</ext-link> describe a novel C-type lectin widely distributed in adult mud crab tissues, <italic>Scylla paramamosain</italic> (<italic>Sp</italic>CTL6), and discuss its immunoprotective effect against bacteria. In turn, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fimmu.2021.679767">Cao et&#xa0;al.</ext-link> report a new lectin expressed by the hepatopancreas of the red swamp crayfish (<italic>Procambarus clarkii</italic>) and discuss its agglutinating ability to promote bacterial clearance.</p>
</sec>
<sec id="s3">
<title>Antiviral Immunity</title>
<p>In penaeid shrimp farming, selective breeding and biosecurity have become essential to prevent viral diseases based on the host&#x2019;s robust resistance to specific pathogens (<xref ref-type="bibr" rid="B2">2</xref>). Both disease prevention and genetic improvement have been based on theoretical support for the interaction between host immunity and virus pathogenesis (<xref ref-type="bibr" rid="B3">3</xref>). In this Research Topic, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fimmu.2020.607543">Zhang et&#xa0;al.</ext-link> report and discuss that glycogen synthase kinase-3&#x3b2; (GSK3&#x3b2;) from shrimp&#xa0;<italic>Litopenaeus vannamei</italic>&#xa0;(<italic>Lv</italic>GSK3&#x3b2;) negatively regulates the activity of NF-&#x3ba;B pathway; RNAi mediated knockdown of&#xa0;<italic>Lv</italic>GSK3&#x3b2; raise the survival of the shrimp infected by White spot syndrome virus (WSSV). In turn, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fimmu.2020.02110">Li et&#xa0;al.</ext-link> reveal that the TRAF3 ortholog from the&#xa0;<italic>L. vannamei&#xa0;</italic>(<italic>Lv</italic>TRAF3) could protect shrimp against WSSV infection by mediating antiviral defense throughout the IRF-Vago pathway, but not by the NF-&#x43a;B pathway. Altogether, these findings describe shrimp immune responses against WSSV and suggest potential candidate genes for resistance-based antiviral defense. Another relevant contribution is made by <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fimmu.2020.01904">Liao et&#xa0;al.</ext-link> concerning the decapod iridescent virus 1 (DIV1), an emergent virus responsible for recent economic troubles in Asian shrimp farming (<xref ref-type="bibr" rid="B4">4</xref>). Comparative transcriptome analysis reveals that DIV1 could hijack the triosephosphate isomerase-like genes (LvTPI) to facilitate virus replication in shrimp (<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fimmu.2020.01904">Liao et&#xa0;al.</ext-link>). Consequently, LvTPI gene silencing reduces shrimp mortality by inhibiting viral replication. This study provides a theoretical basis for the epidemiological investigation of DIV1 infection and may help prevent viral diseases in shrimp farming.</p>
</sec>
<sec id="s4">
<title>Cell-Mediated Responses</title>
<p>Hemocytes represent the immunocompetent cells of crustaceans and mollusks, responsible for cell-mediated responses against invading microorganisms and parasites. Regarding oyster immunity, cell-mediated responses include infiltration, phagocytosis, extracellular trap cell death, encapsulation, immune genes induction, and effectors secretion (<xref ref-type="bibr" rid="B5">5</xref>). In light of this, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fimmu.2021.659469">Lu et&#xa0;al.</ext-link> report and discuss the alteration in some hemato-immunological parameters in oyster <italic>Crassostrea hongkongensis</italic> associated with gender and subpopulation in oysters under bacterial challenge. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fimmu.2021.634497">Estrada et&#xa0;al.</ext-link> evaluated and discuss the occurrence of programmed cell death (PCD), apoptosis and pyroptosis-like, in hemocytes from <italic>Crassostrea gigas</italic> triggered by marine toxins produced by microalgae and bacteria.</p>
</sec>
<sec id="s5">
<title>Host-Pathogen and Microbiota Interactions</title>
<p>Recent studies have revealed exciting aspects of the immune defense triggered by aquatic invertebrates in response to strategies used by pathogens to circumvent them. Insights into host-pathogen and microbiota interactions in fighting invaders have only recently started to emerge. In line with this notion, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fimmu.2021.630343">Petton et&#xa0;al.</ext-link> review and shed light on how the polymicrobial and multifactorial factors operate in the Pacific oyster mortality syndrome (POMS), the most prevalent disease in <italic>C. gigas</italic>. The authors underline the effect of environmental culture conditions and the oyster microbiota imbalance (dysbiosis) in promoting the POMS pathogenesis. Bivalves are sessile filter feeders that live closely associated with large and diverse communities of microorganisms that form their microbiota. In light of this, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fimmu.2020.599625">Gonz&#xe1;lez et&#xa0;al.</ext-link> provide new insights into the interaction between the immune responses of scallop <italic>Argopecten purpuratus</italic> and its microbiota. Scallop antimicrobial peptides and proteins are implicated in maintaining microbial homeostasis and are critical molecules in orchestrating host-microbiota interactions. In turn, aspects of the infection of Chinese crab<italic>, Eriocheir sinensis</italic> by the pathogenic yeast are described by <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fimmu.2021.659723">Jiang et&#xa0;al.</ext-link>, using proteomic analysis. The disease caused by <italic>Metschnikowia bicuspidata</italic> seems to induce some antimicrobial mechanisms (proPO activation system, ROS, and phagocytosis), while the fungal infection suppresses hemolymph coagulation and tissue regeneration.</p>
</sec>
<sec id="s6">
<title>Host Susceptibility Versus Tolerance to Microbial and Viral Infections</title>
<p>As already mentioned, microbial and viral infections have impacted aquatic invertebrates farming, and selective breeding and biosecurity in farms have become essential to prevent diseases based on the host&#x2019;s robust resistance to specific pathogens (<xref ref-type="bibr" rid="B2">2</xref>). Surprisingly, the molecular mechanisms that drive the susceptibility and tolerance of aquatic invertebrates against pathogens remain elusive. A study conducted by <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fimmu.2021.634152">Mai et&#xa0;al.</ext-link> report molecular aspects of differential gene expression in susceptible and resistant strains of shrimp to acute hepatopancreatic necrosis disease (AHPND). A set of candidate genes is proposed to be involved in bacterial pathogenesis, and these findings may aid in breeding programs to select AHPND resistant/tolerant shrimp. In turn, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fimmu.2021.674216">Escobedo-Bonilla</ext-link> provides a broad overview of the viral interference phenomenon in cultivated crustacean species. Exclusion of superinfection, virus-virus interaction, host cell receptor blockade, viral co-infection, and current assays to assess viral interference are widely discussed. Ultimately, a heterologous virus interference mechanism is proposed as a natural strategy to control the incidence of viral diseases in shrimp farming.</p>
</sec>
<sec id="s7">
<title>Concluding Remarks</title>
<p>This Research Topic brings new studies and insights into the molecular, cellular, and physiological mechanisms implicated in antimicrobial and antiviral defenses triggered by the aquatic invertebrates. Original articles and reviews provide novel sharpness into the intricate puzzle of pathogen-host interaction, the role of microbiota in host health and homeostasis, intracellular pathways driving immune effectors/regulators, viral interference phenomena, host susceptibility and tolerance to infections, and other issues. Elucidating the multifaceted mechanisms driving the immunity of farmed aquatic invertebrates is crucial to building new non-antibody-based therapies and controlling aquaculture diseases. Hopefully, the exciting outcomes displayed here can contribute to developing tools that increase shellfish immunocompetence and ultimately promote health security in aquaculture.</p>
</sec>
<sec id="s8" sec-type="author-contributions">
<title>Author Contributions</title>
<p>All authors co-edited the Research Topic. LMP substantially contributed to the conception, drafting, and editing of the final version of the Editorial. CL and KS contributed by drafting part of the work and approved it for publication. All authors contributed to the article and approved the submitted version.</p>
</sec>
<sec id="s9" sec-type="COI-statement">
<title>Conflict of Interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s10" sec-type="disclaimer">
<title>Publisher&#x2019;s Note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
</body>
<back>
<ack>
<title>Acknowledgments</title>
<p>The Editors would like to thank all Research Topic Authors and additional editors for their contributions.</p>
</ack>
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