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<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Plant Sci.</journal-id>
<journal-title>Frontiers in Plant Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Plant Sci.</abbrev-journal-title>
<issn pub-type="epub">1664-462X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fpls.2023.1267183</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Plant Science</subject>
<subj-group>
<subject>Editorial</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Editorial: Revisiting the limits of plant life - plant adaptations to extreme terrestrial environments relating to astrobiology and space biology</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Zupanska</surname><given-names>Agata K.</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/55408"/>
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<contrib contrib-type="author">
<name>
<surname>Arena</surname><given-names>Carmen</given-names>
</name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/124496"/>
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</contrib>
<contrib contrib-type="author">
<name>
<surname>Zu&#xf1;iga</surname><given-names>Gustavo E.</given-names>
</name>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/183805"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
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<contrib contrib-type="author">
<name>
<surname>Casanova-Katny</surname><given-names>Angelica</given-names>
</name>
<xref ref-type="aff" rid="aff4"><sup>4</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/1039707"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
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<contrib contrib-type="author">
<name>
<surname>Turnbull</surname><given-names>Johanna D.</given-names>
</name>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/814200"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Bravo</surname><given-names>Le&#xf3;n A.</given-names>
</name>
<xref ref-type="aff" rid="aff6"><sup>6</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/570570"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ramos</surname><given-names>Patricio</given-names>
</name>
<xref ref-type="aff" rid="aff7"><sup>7</sup></xref>
<xref ref-type="aff" rid="aff8"><sup>8</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/374259"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Sun</surname><given-names>Hang</given-names>
</name>
<xref ref-type="aff" rid="aff9"><sup>9</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/415977"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Shishov</surname><given-names>Vladimir V.</given-names>
</name>
<xref ref-type="aff" rid="aff10"><sup>10</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/584985"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
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</contrib-group>
<aff id="aff1"><sup>1</sup><institution>Carl Sagan Center, SETI Institute</institution>, <addr-line>Mountain View, CA</addr-line>, <country>United States</country></aff>
<aff id="aff2"><sup>2</sup><institution>Department of Biology, University of Naples Federico II</institution>, <addr-line>Naples</addr-line>, <country>Italy</country></aff>
<aff id="aff3"><sup>3</sup><institution>Laboratorio de Fisiolog&#xed;a y Biotecnolog&#xed;a Vegetal, Universidad de Santiago de Chile</institution>, <addr-line>Santiago</addr-line>, <country>Chile</country></aff>
<aff id="aff4"><sup>4</sup><institution>Department of Environmental Sciences, Faculty of Natural Resources, Catholic University of Temuco</institution>, <addr-line>Temuco</addr-line>, <country>Chile</country></aff>
<aff id="aff5"><sup>5</sup><institution>School of Earth, Atmospheric and Life Sciences (SEALS), University of Wollongong</institution>, <addr-line>Wollongong, NSW</addr-line>, <country>Australia</country></aff>
<aff id="aff6"><sup>6</sup><institution>University of La Frontera</institution>, <addr-line>Temuco</addr-line>, <country>Chile</country></aff>
<aff id="aff7"><sup>7</sup><institution>Instituto de Ciencias Biol&#xf3;gicas, Universidad de Talca</institution>, <addr-line>Talca</addr-line>, <country>Chile</country></aff>
<aff id="aff8"><sup>8</sup><institution>Vicerrector&#xed;a de Investigaci&#xf3;n y Postgrado, Universidad Cat&#xf3;lica del Maule</institution>, <addr-line>Talca</addr-line>, <country>Chile</country></aff>
<aff id="aff9"><sup>9</sup><institution>Kunming Institute of Botany, Chinese Academy of Sciences (CAS)</institution>, <addr-line>Kunming</addr-line>, <country>China</country></aff>
<aff id="aff10"><sup>10</sup><institution>School of Fundamental Biology and Biotechnology, Siberian Federal University</institution>, <addr-line>Krasnoyarsk</addr-line>, <country>Russia</country></aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited and Reviewed by: Anna N Stepanova, North Carolina State University, United States</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Agata Klaudia Zupanska, <email xlink:href="mailto:azupanska@seti.org">azupanska@seti.org</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>17</day>
<month>08</month>
<year>2023</year>
</pub-date>
<pub-date pub-type="collection">
<year>2023</year>
</pub-date>
<volume>14</volume>
<elocation-id>1267183</elocation-id>
<history>
<date date-type="received">
<day>26</day>
<month>07</month>
<year>2023</year>
</date>
<date date-type="accepted">
<day>07</day>
<month>08</month>
<year>2023</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2023 Zupanska, Arena, Zu&#xf1;iga, Casanova-Katny, Turnbull, Bravo, Ramos, Sun and Shishov</copyright-statement>
<copyright-year>2023</copyright-year>
<copyright-holder>Zupanska, Arena, Zu&#xf1;iga, Casanova-Katny, Turnbull, Bravo, Ramos, Sun and Shishov</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/46160" ext-link-type="uri">Editorial on the Research Topic <article-title>Revisiting the limits of plant life - plant adaptations to extreme terrestrial environments relating to astrobiology and space biology</article-title>
</related-article>
<kwd-group>
<kwd>plant</kwd>
<kwd>extreme environment</kwd>
<kwd>astrobiology</kwd>
<kwd>deep space exploration</kwd>
<kwd>limits of life</kwd>
</kwd-group>
<counts>
<fig-count count="0"/>
<table-count count="0"/>
<equation-count count="0"/>
<ref-count count="7"/>
<page-count count="3"/>
<word-count count="1088"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Plant Physiology</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<p>Plants were essential to the early evolution of terrestrial life and colonization of the young Earth (<xref ref-type="bibr" rid="B5">Kapoor et&#xa0;al., 2023</xref>). Plant communities continue to colonize and transform our planet including the newest ecosystems formed post-glaciation, restoring those degraded by human activities and adapting to changing ecological conditions (<xref ref-type="bibr" rid="B3">Huston and Smith, 1987</xref>; <xref ref-type="bibr" rid="B1">Chapin et&#xa0;al., 1994</xref>; <xref ref-type="bibr" rid="B7">Yuan et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B2">Heim et&#xa0;al., 2021</xref>). Plants cannot move away from a harmful stimulus, and thus, have evolved remarkable strategies to survive and eventually thrive in harsh environments. Today, humanity is on the verge of exploring our solar system and beyond, eager to discover, answer fundamental questions, and search for extraterrestrial forms of life. Undoubtedly, plants are key organisms to successful deep space missions and independence from the provision of terrestrial resources, whether for long duration interplanetary travel or establishing permanent settlements. We can employ terrestrial bio-design principles to mimic plant colonization on Earth and explore adaptations to novel ecosystems. This can be combined with precise molecular tools to enable long-term human space exploration. With this thought in mind, we have collated articles focusing on terrestrial plants from extreme environments and their adaptations to harsh conditions. This collective knowledge will advance the selection of desired plant characteristics relevant to human space missions and assist in the identification of valuable adaptive mechanisms that can be genetically engineered in targeted space food plants. Furthermore, terrestrial plants from the edge of habitability define limits for terrestrial forms of life and thus can direct efforts for the search for life beyond Earth (<xref ref-type="bibr" rid="B6">McKay, 2014</xref>; <xref ref-type="bibr" rid="B4">Huwe et&#xa0;al., 2019</xref>).</p>
<p>In this Research Topic, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2022.1001158">De Micco et&#xa0;al.</ext-link> present an exhaustive review on the hazards of cosmic ionizing radiation to human exploration and discuss potential roles for plants in mitigating this Research Topic. The authors highlight difficulties in studying ionizing radiation in deep space that stem from deploying inadequate radiation sources, limited access to ion beam accelerators, and lack of uniformity in radiation doses and dose rates (acute, chronic) across studies. The authors expand on the remarkable resistance of plants growing in regions of high radioactivity and the possibility of using such plants in shielding or for pharmaceutical countermeasures. Next, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1139704">Molina-Montenegro et&#xa0;al.</ext-link> demonstrate the benefits of co-culturing relevant crops with endophytic fungi in simulated deep space environments to augment plant performance in high UV radiation, low temperature, and low water availability. In this instance, plants from mesic habitats were combined with fungi from the extreme environment of the Atacama Desert, which as the driest place in the world and also experiences excessive solar irradiance. Increased survival in crops inoculated with desert fungi was related to higher biomass, antioxidant content and nutritional quality in deep space-like environmental conditions compared to non-inoculated plants. Thus, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1139704">Molina- Montenegro et&#xa0;al.</ext-link> contribute to developing the use of symbionts from extreme terrestrial environments to improve the efficiency of crop cultivation in space.</p>
<p>In another study, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1110521">Gong et&#xa0;al.</ext-link> challenged emerging seedlings with growth mediums of varying particle size, porosity, and compactness, which is pivotal to understanding the intricacies of germinating seeds in extraterrestrial regolith in future space farms. Extraterrestrial regolith on anhydrous and airless celestial bodies is formed by vastly different processes than terrestrial soils. These processes include large and small meteoroid impacts, star plasma winds, and cosmic ionizing radiation, and cause distinct mechanical, physical, and hydraulic properties. Gravitational effects further impact regolith formation by affecting collision and aggregation velocities, which influences the values of bulk densities and the porosity of the outermost regolith layers. Using CT scans corroborated by mathematical models, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1110521">Gong et&#xa0;al.</ext-link> demonstrate that underground seedlings search for a path of least resistance irrespective of the distance to the surface, and that high porosity and low compactness of the growth medium are conducive to seedling emergence. This finding informs plant cultivation on the Moon using <italic>in situ</italic> regolith. Due to micrometeoroid impacts, lunar regolith has very fine, low- density particles with complex shapes, sharp jagged edges, and a high angular surface. According to <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1110521">Gong et&#xa0;al.</ext-link>, these tight and abrasive pore spaces could impede seedling path to the surface. However, the lower gravitational field on the Moon results in much lower regolith bulk densities, hence low compactness, which, according to the findings of <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1110521">Gong et&#xa0;al.</ext-link>, is likely to promote seedling emergence.</p>
<p>Another study in this Research Topic addressed the quest for extraterrestrial life. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1070359">Battistuzzi et&#xa0;al.</ext-link> explore if the light spectrum emissions from faint and cool stars is sufficient for oxygenic photosynthesis in cyanobacteria. This is relevant to the search for oxygen biosignatures on exoplanets orbiting M-dwarf or Red Dwarf stars in habitable zones. These stars emit light at the far-red (700&#x2013;750nm) and infrared (750&#x2013;1000nm) wavelengths, unlike the Sun, which emits light predominantly in the visible and UV spectrum. On Earth, photosynthesizing plants harvest solar blue and red light for the light reactions of photosynthesis, so the capacity for low energy photons at longer wavelengths to photolyze H<sub>2</sub>O was unknown. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1070359">Battistuzzi et&#xa0;al.</ext-link> show that cyanobacteria can acclimate to the simulated M-dwarf light spectrum by elevating the concentration of pigments capable of quenching low-energy photons and by increasing the efficiency of utilizing the sparse high-energy photons. There was no difference in biomass production nor O<sub>2</sub> release between cyanobacteria grown under either light spectrum. This demonstrates that the photosynthetic apparatus, which evolved in the earliest terrestrial life forms, could perform in the presence of cool and dim starlight. As such, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1070359">Battistuzzi et&#xa0;al.</ext-link> contribute a rare physiological analysis to theoretical models evaluating the feasibility of M-dwarf stars to support photosynthesis. This informs and legitimizes the search for oxygen biosignatures.</p>
<p>This Research Topic was conceived with the intent to present research, methods, and reviews to further our understanding of plant survival in harsh or novel environments, and that can be extrapolated to the conditions found in deep space. As humanity leaves Earth&#x2019;s protective influence to explore the solar system and beyond, we require the knowledge and ideas developed by plant scientists, such as those assembled in this Research Topic, to take plants with us and deploy them to perform mission-critical functions. Once at the destination, guided by the knowledge of plant evolutionary solutions, we will know where to look for extraterrestrial life.</p>
<sec id="s1" sec-type="author-contributions">
<title>Author contributions</title>
<p>AZ: Writing &#x2013; original draft. CA: Writing &#x2013; review &amp; editing, Data curation. GZ: Writing &#x2013; review &amp; editing. AC-K: Writing &#x2013; review &amp; editing. JT: Writing &#x2013; review &amp; editing. LB: Writing &#x2013; review &amp; editing. PR: Writing &#x2013; review &amp; editing. HS: Writing &#x2013; review &amp; editing. VS: Writing &#x2013; review &amp; editing.</p>
</sec>
</body>
<back>
<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="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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