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<front>
<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>
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<article-meta>
<article-id pub-id-type="doi">10.3389/fpls.2022.1126610</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: Ecoepigenetics in clonal and inbreeding plants: Transgenerational adaptation and environmental variation, Volume II</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Dong</surname>
<given-names>Bi-Cheng</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/294618"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Roiloa</surname>
<given-names>Sergio R.</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/204855"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Xue</surname>
<given-names>Wei</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1080127"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Yu</surname>
<given-names>Fei-Hai</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/204452"/>
</contrib>
</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>School of Ecology and Nature Conservation, Beijing Forestry University</institution>, <addr-line>Beijing</addr-line>, <country>China</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>The Key Laboratory of Ecological Protection in the Yellow River Basin of National Forestry and Grassland Administration, Beijing Forestry University</institution>, <addr-line>Beijing</addr-line>, <country>China</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>BioCost Group, Department of Biology, Faculty of Science, University of A Coru&#xf1;a</institution>, <addr-line>A Coru&#xf1;a</addr-line>, <country>Spain</country>
</aff>
<aff id="aff4">
<sup>4</sup>
<institution>Institute of Wetland Ecology &amp; Clone Ecology, Taizhou University</institution>, <addr-line>Taizhou</addr-line>, <country>China</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited and Reviewed by: Xiaohua Jin, Institute of Botany (CAS), China</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Bi-Cheng Dong, <email xlink:href="mailto:bcdong@bjfu.edu.cn">bcdong@bjfu.edu.cn</email>; Fei-Hai Yu, <email xlink:href="mailto:feihiayu@126.com">feihiayu@126.com</email>
</p>
</fn>
<fn fn-type="other" id="fn002">
<p>This article was submitted to Plant Systematics and Evolution, a section of the journal Frontiers in Plant Science</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>11</day>
<month>01</month>
<year>2023</year>
</pub-date>
<pub-date pub-type="collection">
<year>2022</year>
</pub-date>
<volume>13</volume>
<elocation-id>1126610</elocation-id>
<history>
<date date-type="received">
<day>18</day>
<month>12</month>
<year>2022</year>
</date>
<date date-type="accepted">
<day>28</day>
<month>12</month>
<year>2022</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2023 Dong, Roiloa, Xue and Yu</copyright-statement>
<copyright-year>2023</copyright-year>
<copyright-holder>Dong, Roiloa, Xue and Yu</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/28335" ext-link-type="uri">The Editorial on the Research Topic <article-title>Ecoepigenetics in clonal and inbreeding plants: Transgenerational adaptation and environmental variation, Volume II</article-title>
</related-article>
<kwd-group>
<kwd>clonal growth</kwd>
<kwd>environmental change</kwd>
<kwd>epigenetics</kwd>
<kwd>parental effect</kwd>
<kwd>resource provisioning</kwd>
<kwd>transgenerational plasticity</kwd>
<kwd>inbreeding plants</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="1133"/>
</counts>
</article-meta>
</front>
<body>    <p>Accelerating environmental changes at the local, regional, and global scales are likely to favor species that can rapidly adapt to new environmental conditions. Long-lived clonal plants, in which reproduction is mainly asexual and clonal growth plays a central role in their population spread and maintenance, have long been thought to possess low genetic variation so that the potential for genetics-based adaptation to environmental changes may be limited. Thus, epigenetic variation may be particularly important for these clonal plants to adapt to rapid environmental changes (<xref ref-type="bibr" rid="B6">Latzel and Klime&#x161;ov&#xe1;, 2010</xref>; <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fevo.2016.00086">Dodd and Douhovnikoff</ext-link>; <xref ref-type="bibr" rid="B8">Mounger et&#xa0;al., 2021</xref>).</p>
<p>Recent work suggests that the performance of an individual (ramet) of clonal plants is influenced by not only its current environmental condition but also the environmental condition of its parents (<xref ref-type="bibr" rid="B5">Huber et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B9">Xue et&#xa0;al., 2022</xref>). At least three mechanisms can explain such transgenerational (parental or maternal) effects (<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2011.00102">Herman and Sultan</ext-link>; <xref ref-type="bibr" rid="B7">Luo et&#xa0;al., 2022</xref>). First, parental environments could directly influence the performance of clonal offspring by altering the provisioning of carbohydrates and nutrients in vegetative propagules (e.g., fragmented stolons, rhizomes, or storage roots) (<xref ref-type="bibr" rid="B2">Dong et&#xa0;al., 2019</xref>). Second, environmental stress could also induce non-provisioning effects between clonal generations, <italic>via</italic> modifying the allocation of defensive chemicals and/or defence-inducing hormones to clonal offspring (<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2011.00102">Herman and Sultan</ext-link>). Third, parental environments could trigger epigenetic changes in parent plants (e.g., the methylation of DNA and modifications of histones), to facilitate and optimize phenotype variation of clonal offspring in response to environmental change (<xref ref-type="bibr" rid="B3">Douhovnikoff and Dodd, 2015</xref>). This Research Topic consists of 11 articles, most of which explore the ecological significance of transgenerational effects and epigenetic variation in clonal plants.</p>
<p>Three papers focus on the relationship between epigenetic regulation and local adaptation of clonal plants under variable environmental stress. By experimental demethylation in natural conditions across different regions of Europe, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2022.827166">Sammarco et&#xa0;al.</ext-link> found that the local adaptation mediated by epigenetic variation allowed the stoloniferous plant <italic>Fragaria vesca</italic> to better respond to changing climatic conditions. They suggest that epigenetic-based local adaptation may provide clonal plants with sufficient time to tackle the ongoing environmental crisis and to genetically adapt to it afterwards. In a field experiment, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2022.888391">Campoy et&#xa0;al.</ext-link> compared variations in DNA methylation and phenotypic traits between native and introduced populations of the clonal succulent species <italic>Carpobrotus edulis</italic> under a climate change scenario, showing that phenotypic plasticity and global DNA methylation might be related to its rapid adaptation to new habitats. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2022.831175">Wang et&#xa0;al.</ext-link> grew experimental populations of the creeping plant <italic>Hydrocotyle vulgaris</italic>, consisting of the same genotype, in two flood regimes and found significant phenotypic differences and associated DNA methylation differentiation between the two types of populations. They suggest that DNA methylation was involved in plant responses to environmental variation.</p>
<p>Four papers consider clonal transgenerational effects on growth, stress tolerance, and competitive ability of clonal offspring. Calibrated with data from two experiments, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2022.872065">Wang et&#xa0;al.</ext-link> developed a model to test the transgenerational nitrogen effects on the summed and the mean performance of clonal offspring of the creeping clonal plant <italic>Alternanthera philoxeroides</italic>. They found that transgenerational effects at the whole-generation scale could be jointly influenced by multiple plant inherent characteristics (e.g., the survival rate, the number and the size distribution of clonal propagules), and the magnitude of transgenerational effects could also be obscured by developmental constraints. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2022.890309">Zhang et&#xa0;al.</ext-link> tested transgenerational nitrogen effects on the fitness of three generations of the floating clonal plant <italic>Pistia stratiotes</italic>. They found that resource provisioning can increase the initial establishment of clonal offspring in favourable conditions, but this effect may not always be beneficial to their subsequent growth. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2022.924001">Yu et&#xa0;al.</ext-link> showed that transgenerational effects could regulate interspecific competition between <italic>P. stratiotes</italic> and <italic>Eichhornia crassipes</italic> by altering the competitive ability of <italic>P. stratiotes</italic>, <italic>via</italic> changes in resource provisioning and/or DNA methylation. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2022.949752">Guo et&#xa0;al</ext-link>. tested transgenerational ultraviolet-B (UV-B) effects on the fitness of clonal offspring of the stoloniferous plant <italic>Glechoma longituba</italic>. They found that transgenerational effects could promote the increase in the biomass allocation to aboveground parts in clonal offspring under similar UV-B stress, as well as their defence substances (e.g., flavonoid and anthocyanin), suggesting that the anticipatory transgenerational effects were likely to improve the UV-B resistance.</p>
<p>Two papers examine the effects of population differentiation on the offspring performance of widely-distributed species. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2022.919409">Chen et&#xa0;al.</ext-link> examined whether parental environments (i.e., plants were collected from the high and low elevations in the hydro-fluctuation belt of the <italic>Three Gorges Reservoir</italic> region) and the early exposure of offspring of <italic>Polygonum hydropiper</italic> to flooding (accompanied with or without eutrophication) became as a positive or stressful cue on the subsequent growth of these offspring. They found that offspring produced by parental plants in the low elevation might have high adaptability in response to this &#x201c;predictable&#x201d; periodic flooding stress. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2022.994367">Liu et&#xa0;al</ext-link>. tested the effects of populations with different introduction histories on growth traits of an invasive herb <italic>Erigeron annuus</italic> both in the wild and in common garden experiments. They found that there was parallel genetic and phenotypic differentiation among different invasive populations and that the populations that were introduced earlier had higher genetic diversity and higher growth dominance.</p>
<p>Two papers report within-generation responses of clonal species to stressful environments. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2022.831654">Qi et&#xa0;al.</ext-link> tested the interaction effects between arbuscular mycorrhizal fungi (AMF) and soil phosphorus availability on the uptake ability and allocation strategies of an invasive clonal herb <italic>Solidago canadensis</italic>. They found that AMF were able to facilitate phosphorous acquisition by <italic>S. canadensis</italic> in insoluble phosphorous conditions, and also contribute to the invasiveness of <italic>S. canadensis</italic> in the resource-deficient environment. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2022.883800">Jing et&#xa0;al.</ext-link> tested the effects of submergence depths on the growth responses of the clonal herb <italic>A. philoxeroides</italic>. They found that <italic>A. philoxeroides</italic> switched from &#x201c;escape&#x201d; to &#x201c;quiescence&#x201d; strategies in response to increasing submergence depths, and that morphological plasticity such as stem elongation could be essential for the acclimatization of <italic>A. philoxeroides</italic> to water-level fluctuations.</p>
<p>Transgenerational effects in clonal plants have drawn increasing attention during the last few years (<xref ref-type="bibr" rid="B7">Luo et&#xa0;al., 2022</xref>). However, there is still a long way to explore in this exciting field. Thus, the knowledge of the mechanisms that relate transgenerational environmental effects to epigenetic inheritance in clonal plants, the correlation between epigenetic variation with genetic and phenotypic variation in wild plant populations, the ecological and evolutionary role of transgenerational effects at different scales (e.g., the individual, population and community levels) have rarely been investigated so far. With the publications on this topic, we hope to further fill in the knowledge gap and stimulate more research on this important issue in the future.</p>
<sec id="s1" sec-type="author-contributions">
<title>Author contributions</title>
<p>All authors listed have made substantial and direct intellectual contributions to the work and approved it for publication.</p>
</sec>
</body>
<back>
<sec id="s2" sec-type="funding-information">
<title>Funding</title>
<p>Research was supported by grants from the National Natural Science Foundation of China (32071527) to F-HY, and the National Natural Science Foundation of China (31500331) to B-CD.</p>
</sec>
<sec id="s3" 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="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>
<ref-list>
<title>References</title>
<ref id="B2">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Dong</surname> <given-names>B.-C.</given-names>
</name>
<name>
<surname>Alpert</surname> <given-names>P.</given-names>
</name>
<name>
<surname>Yu</surname> <given-names>F.-H.</given-names>
</name>
</person-group> (<year>2019</year>). <article-title>Transgenerational effects of herbivory and soil nutrients transmitted <italic>via</italic> vegetative reproduction in the clonal plant <italic>Alternanthera philoxeroides</italic>
</article-title>. <source>Perspect. Plant Ecol. Evol. Syst.</source> <volume>41</volume>, <elocation-id>125498</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.ppees.2019.125498</pub-id>
</citation>
</ref>
<ref id="B3">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Douhovnikoff</surname> <given-names>V.</given-names>
</name>
<name>
<surname>Dodd</surname> <given-names>R. S.</given-names>
</name>
</person-group> (<year>2015</year>). <article-title>Epigenetics: A potential mechanism for clonal plant success</article-title>. <source>Plant Ecol.</source> <volume>216</volume>, <fpage>227</fpage>&#x2013;<lpage>233</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s11258-014-0430-z</pub-id>
</citation>
</ref>
<ref id="B5">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Huber</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Gablenz</surname> <given-names>S.</given-names>
</name>
<name>
<surname>H&#xf6;fer</surname> <given-names>M.</given-names>
</name>
</person-group> (<year>2021</year>). <article-title>Transgenerational non-genetic inheritance has fitness costs and benefits under recurring stress in the clonal duckweed <italic>Spirodela polyrhiza</italic>
</article-title>. <source>Proc. R. Soc B Biol. Sci.</source> <volume>288</volume>, <fpage>20211269</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1098/rspb.2021.1269</pub-id>
</citation>
</ref>
<ref id="B6">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Latzel</surname> <given-names>V.</given-names>
</name>
<name>
<surname>Klime&#x161;ov&#xe1;</surname> <given-names>J.</given-names>
</name>
</person-group> (<year>2010</year>). <article-title>Transgenerational plasticity in clonal plants</article-title>. <source>Evol. Ecol.</source> <volume>24</volume>, <fpage>1537</fpage>&#x2013;<lpage>1543</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s10682-010-9385-2</pub-id>
</citation>
</ref>
<ref id="B7">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Luo</surname> <given-names>F.-L.</given-names>
</name>
<name>
<surname>Wang</surname> <given-names>M.-Z.</given-names>
</name>
<name>
<surname>Dong</surname> <given-names>B.-C.</given-names>
</name>
<name>
<surname>Chen</surname> <given-names>Y.-H.</given-names>
</name>
<name>
<surname>Yu</surname> <given-names>F.-H.</given-names>
</name>
</person-group> (<year>2022</year>). <article-title>Clonal and sexual parental effects and their mechanisms</article-title> (in Chinese). <source>Acta Ecol. Sin.</source> <volume>42</volume>, <fpage>6474</fpage>&#x2013;<lpage>6486</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.5846/stxb202107211968</pub-id>
</citation>
</ref>
<ref id="B8">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Mounger</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Ainouche</surname> <given-names>M. L.</given-names>
</name>
<name>
<surname>Bossdorf</surname> <given-names>O.</given-names>
</name>
<name>
<surname>Cav&#xe9;-Radet</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Li</surname> <given-names>B.</given-names>
</name>
<name>
<surname>Parepa</surname> <given-names>M.</given-names>
</name>
<etal/>
</person-group>. (<year>2021</year>). <article-title>Epigenetics and the success of invasive plants</article-title>. <source>Philos. Trans. R. Soc B Biol. Sci.</source> <volume>376</volume>, <fpage>20200117</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1098/rstb.2020.0117</pub-id>
</citation>
</ref>
<ref id="B9">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Xue</surname> <given-names>W.</given-names>
</name>
<name>
<surname>Huang</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Yu</surname> <given-names>F.-H.</given-names>
</name>
<name>
<surname>Bezemer</surname> <given-names>T. M.</given-names>
</name>
</person-group> (<year>2022</year>). <article-title>Light condition experienced by parent plants influences the response of offspring to light <italic>via</italic> both parental effects and soil legacy effects</article-title>. <source>Funct. Ecol.</source> <volume>36</volume>, <fpage>2434</fpage>&#x2013;<lpage>2444</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/1365-2435.14136</pub-id>
</citation>
</ref>
</ref-list>
</back>
</article>