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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Ecol. Evol.</journal-id>
<journal-title>Frontiers in Ecology and Evolution</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Ecol. Evol.</abbrev-journal-title>
<issn pub-type="epub">2296-701X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fevo.2025.1609597</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Ecology and Evolution</subject>
<subj-group>
<subject>Editorial</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Editorial: Biological and physical basis of the development of integument and associated structures</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Finet</surname>
<given-names>C&#xe9;dric</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/246024/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
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<contrib contrib-type="author" corresp="yes">
<name>
<surname>Prakash</surname>
<given-names>Anupama</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<xref ref-type="author-notes" rid="fn003">
<sup>&#x2020;</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/553702/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
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<aff id="aff1">
<sup>1</sup>
<institution>Department of Biological Sciences, National University of Singapore</institution>, <addr-line>Singapore</addr-line>, <country>Singapore</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Ecology and Evolutionary Biology, School of Biosciences, University of Sheffield</institution>, <addr-line>Sheffield</addr-line>, <country>United Kingdom</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited and Reviewed by: Maria Ina Arnone, Stazione Zoologica Anton Dohrn, Italy</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: C&#xe9;dric Finet, <email xlink:href="mailto:cedric.finet@ens-lyon.org">cedric.finet@ens-lyon.org</email>; Anupama Prakash, <email xlink:href="mailto:a.sarojini-prakash@imperial.ac.uk">a.sarojini-prakash@imperial.ac.uk</email>
</p>
</fn>
<fn fn-type="present-address" id="fn003">
<p>&#x2020;Present address: Anupama Prakash, Department of Bioengineering, Imperial College, London, United Kingdom</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>06</day>
<month>05</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>13</volume>
<elocation-id>1609597</elocation-id>
<history>
<date date-type="received">
<day>10</day>
<month>04</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>22</day>
<month>04</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2025 Finet and Prakash</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Finet and Prakash</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" journal-id="Front Ecol Evol" journal-id-type="nlm-ta" xlink:href="https://www.frontiersin.org/research-topics/54743/biological-and-physical-basis-of-the-development-of-integument-and-associated-structures" ext-link-type="uri">Editorial on the Research Topic <article-title>Biological and physical basis of the development of integument and associated structures</article-title>
</related-article>
<kwd-group>
<kwd>integument</kwd>
<kwd>micro- and nano-structures</kwd>
<kwd>morphogenesis</kwd>
<kwd>biomaterials</kwd>
<kwd>bioinspiration and biomimetics</kwd>
</kwd-group>
<counts>
<fig-count count="0"/>
<table-count count="0"/>
<equation-count count="0"/>
<ref-count count="55"/>
<page-count count="4"/>
<word-count count="1063"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Evolutionary Developmental Biology</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<disp-quote>
<p>&#x201c;<italic>Everything about microscopic life is terribly upsetting. How can things so small be so important</italic>?&#x201d;</p>
<p>Isaac Asimov</p>
</disp-quote>
<p>The integument of organisms differs greatly in nature and shape, including the cell wall in bacteria, fungi, algae and plants, the cuticle in arthropods, and the skin in vertebrates. The integumentary surface offers a range of micro- and nano-structures that serve a variety of purposes like environmental sensing, light scattering, substrate adhesion, (super)hydrophobicity, (super)hydrophilicity, and thermoregulation (<xref ref-type="bibr" rid="B6">Barthlott et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B53">Watson et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B46">Seale et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B3">Akat et&#xa0;al., 2022</xref>). It is worth noting that these structures are often multifunctional. Water striders, for instance, have leg bristles that give both mechanosensation and water repellency (<xref ref-type="bibr" rid="B14">Finet et&#xa0;al., 2018</xref>, <xref ref-type="bibr" rid="B15">2022</xref>), whilst clearwing butterflies have transparent wings with anti-reflective and hydrophobic nipple arrays (<xref ref-type="bibr" rid="B16">Finet et&#xa0;al., 2023</xref>).</p>
<p>While integuments and their accompanying structures are extremely diverse, their material composition is the result of developmental and evolutionary tinkering of a small set of biopolymers like chitin, keratin, and cellulose, as well as various proteins, lipids, and pigments (<xref ref-type="bibr" rid="B39">Pasquina-Lemonche et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B3">Akat et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B37">Muthukrishnan et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B20">Gow and Lenardon, 2023</xref>; <xref ref-type="bibr" rid="B10">Cosgrove, 2024</xref>). Structural colors for example, can be created by combinations of these biomaterials (<xref ref-type="bibr" rid="B33">McPhedran et&#xa0;al., 2001</xref>; <xref ref-type="bibr" rid="B45">Seago et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B47">Sun and Bhushan, 2012</xref>; <xref ref-type="bibr" rid="B2">Airoldi et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B22">Hsiung et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B34">Middleton et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B44">Saranathan and Finet, 2021</xref>; <xref ref-type="bibr" rid="B51">Thayer and Patel, 2023</xref>). Furthermore, these biomaterials are optimized and often hierarchically structured, indicating precise cellular/tissue control over biomaterial assembly across space and time (<xref ref-type="bibr" rid="B35">Miserez et&#xa0;al., 2008</xref>; <xref ref-type="bibr" rid="B8">Carroll et&#xa0;al., 2022</xref>).</p>
<p>The morphogenesis of integumentary surfaces and nanostructures remains a vast and underexplored field. However, the emergence of recent reviews (<xref ref-type="bibr" rid="B2">Airoldi et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B29">Lloyd and Nadeau, 2021</xref>; <xref ref-type="bibr" rid="B44">Saranathan and Finet, 2021</xref>; <xref ref-type="bibr" rid="B13">Finet, 2024</xref>) and research papers on this topic highlights an active field of research. Our understanding of the spatial control of chitin assembly at the subcellular level has progressed (<xref ref-type="bibr" rid="B40">Pesch et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B1">Adler, 2019</xref>; <xref ref-type="bibr" rid="B49">Sviben et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B11">De Giorgio et&#xa0;al., 2023</xref>; <xref ref-type="bibr" rid="B19">Ghosh and Treisman, 2024</xref>; <xref ref-type="bibr" rid="B23">Ikeda et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B24">Inagaki et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B25">Itakura et&#xa0;al., 2024</xref>). Single-cell gene expression atlases of developing scales in butterflies (<xref ref-type="bibr" rid="B41">Prakash et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B30">Loh et&#xa0;al., 2025</xref>) and bristles in <italic>Drosophila</italic> (<xref ref-type="bibr" rid="B21">Hopkins et&#xa0;al., 2023</xref>) have identified gene networks involved in hair-like structure morphogenesis. Progress has been made to understand the formation of different nanostructures in butterfly wing scales such as the laminae (<xref ref-type="bibr" rid="B50">Thayer et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B42">Prakash et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B9">Chatterjee et&#xa0;al., 2023</xref>; <xref ref-type="bibr" rid="B4">Balakrishnan et&#xa0;al., 2024</xref>), the ridges (<xref ref-type="bibr" rid="B7">Brien et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B12">Ficarrotta et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B28">Lloyd et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B52">Totz et&#xa0;al., 2024</xref>), the luminal gyroid (<xref ref-type="bibr" rid="B54">Wilts et&#xa0;al., 2017</xref>), and the trabeculae (<xref ref-type="bibr" rid="B43">Ru et&#xa0;al., 2024</xref>). Work on cuticular proteins has identified their roles and spatial distributions in the development of beetle elytra (<xref ref-type="bibr" rid="B38">Noh et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B36">Murata et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B5">Bao et&#xa0;al., 2024</xref>) and butterfly scales (<xref ref-type="bibr" rid="B27">Liu et&#xa0;al., 2021</xref>).</p>
<p>In particular, many studies feed into the biomimetic and bioengineering fields of research. For example, the natural world has inspired multiple solutions for material adhesion (<xref ref-type="bibr" rid="B26">Li et&#xa0;al., 2024</xref>). The morphology of shark scales has led to various applications from membrane antifouling to hydrodynamics (<xref ref-type="bibr" rid="B18">Ghimire et&#xa0;al., 2024</xref>), and chameleons and cephalopods have inspired color-changing hydrogels for multiple functions including sensors (<xref ref-type="bibr" rid="B31">Lu et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B55">Zhang et&#xa0;al., 2023</xref>). However, beyond extracting design principles from biological materials, the ultimate goal would be to develop bio-inspired manufacturing processes, which require an in-depth understanding of the biological processes themselves.</p>
<p>This topic explores the genetics and cellular mechanisms underlying the development of integument and associated structures in animals and plants. With this Research Topic, we wish to direct readers to the emerging field of bio-inspired manufacturing and hope to acknowledge the need for more studies in understanding biological processes that can produce precise and compositionally driven micro- and nano-structures.</p>
<p>Plant cell walls are made up of cellulose microfibrils embedded in a matrix of glycoproteins, and pectic and hemicellulosic polysaccharides. Glycosylphosphatidylinositol (GPI) is a common eukaryotic lipid modification that helps proteins adhere to the membrane lipid bilayer. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fcell.2022.904714">Zhou</ext-link> reviews our knowledge on GPI-anchored proteins involved in cell wall regulation in the plant model <italic>Arabidopsis</italic>. The author proposes that these proteins might act as structural components of the cell wall by organizing cellulose microfibrils at the cell surface.</p>
<p>In reptiles, a mutation in the <italic>TFEC</italic> gene leads to a piebald phenotype with white patches in the ball python while it causes reduced coloration in the brown anole lizard due to the loss of iridophores (<xref ref-type="bibr" rid="B17">Garcia-Elfring et&#xa0;al., 2023</xref>). Using comparative histology, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fcell.2024.1358828">Tzika</ext-link> demonstrates that <italic>TFEC</italic> mutants produce similar phenotypes of reduced coloration via different mechanisms. In the anole, TFEC is necessary for the development of iridophores, whereas in the ball python, which lacks iridophores, TFEC is important for the development of melanophores and xanthophores. By pointing out that the same transcription factor can function differently within the same taxon, this study emphasizes that the phenotypic mutant approach is insufficient for elucidating the underlying molecular mechanisms.</p>
<p>Like reptiles, the ribbontail stingray exhibits structurally colored blue spots produced by dermal iridophores. These iridophores are unique by having numerous fingerlike protuberances (<xref ref-type="bibr" rid="B48">Surapaneni et&#xa0;al., 2024</xref>) and contain spherical iridosomes enclosing guanine nanocrystals. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fcell.2024.1393237">Blumer et&#xa0;al.</ext-link> provide a detailed ultrastructural description of the ribbontail ray&#x2019;s novel iridophore. They found that intermediate filaments form an intracellular scaffold that spaces the iridosomes within the iridophores.</p>
<p>In crickets and grasshoppers, males have evolved cuticular structures on their forewings to produce sound via stridulation. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fevo.2024.1411228">Turchyn and Popadi&#x107;</ext-link> identify the POU homeodomain gene <italic>nubbin</italic> as a regulator of the development of sound resonators on the wings of the house cricket. They propose that <italic>nubbin</italic>, a key player in the wing development network, has been recruited in the course of evolution of Orthoptera to evolve these new cuticular nanostructures.</p>
<p>
<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fevo.2024.1392050">Banerjee et&#xa0;al.</ext-link> investigate the interplay between nanomorphology and pigmentation during the development of butterfly scales. They show that the loss-of-function mutations in <italic>Optix</italic> result in both pigmentation and nanomorphology defects. By comparing these effects with mutants in melanin and/or ommochrome pathways, they propose that Optix regulates nanomorphology via its effects on pigmentation, complementing earlier studies on melanized (<xref ref-type="bibr" rid="B32">Matsuoka and Monteiro, 2018</xref>) and silver scales (<xref ref-type="bibr" rid="B42">Prakash et&#xa0;al., 2022</xref>).</p>
</body>
<back>
<sec id="s1" sec-type="author-contributions">
<title>Author contributions</title>
<p>CF: Writing &#x2013; original draft, Writing &#x2013; review &amp; editing. AP: Writing &#x2013; original draft, Writing &#x2013; review &amp; editing.</p>
</sec>
<sec id="s2" 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="s3" sec-type="ai-statement">
<title>Generative AI statement</title>
<p>The author(s) declare that no Generative AI was used in the creation of this manuscript.</p>
</sec>
<sec id="s4" 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>
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