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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>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fpls.2024.1508507</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: Research advances on forest tree functional genomics and breeding</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Du</surname>
<given-names>Juan</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>
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<contrib contrib-type="author">
<name>
<surname>Ye</surname>
<given-names>Tianqi</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/data-curation/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>An</surname>
<given-names>Yi</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1137633"/>
<role content-type="https://credit.niso.org/contributor-roles/validation/"/>
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</contrib>
<contrib contrib-type="author">
<name>
<surname>Chen</surname>
<given-names>Yicun</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/363029"/>
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<contrib contrib-type="author">
<name>
<surname>Wang</surname>
<given-names>Jack</given-names>
</name>
<xref ref-type="aff" rid="aff5">
<sup>5</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/408657"/>
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<contrib contrib-type="author">
<name>
<surname>Wang</surname>
<given-names>Jiehua</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<xref ref-type="aff" rid="aff6">
<sup>6</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/494349"/>
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</contrib>
<contrib contrib-type="author">
<name>
<surname>Lu</surname>
<given-names>Mengzhu</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/215079"/>
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</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Li</surname>
<given-names>Quanzi</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/660764"/>
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</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>State Key Laboratory of Plant Physiology and Biochemistry, College of Life Sciences, Zhejiang University</institution>, <addr-line>Hangzhou, Zhejiang</addr-line>, <country>China</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Institute of Fundamental and Transdisciplinary, Zhejiang University</institution>, <addr-line>Hangzhou, Zhejiang</addr-line>, <country>China</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>College of Forestry and Biotechnology, Zhejiang A &amp; F University</institution>, <addr-line>Hangzhou</addr-line>, <country>China</country>
</aff>
<aff id="aff4">
<sup>4</sup>
<institution>State Key Laboratory of Tree Genetics and Breeding, Chinese Academy of Forestry</institution>, <addr-line>Beijing</addr-line>, <country>China</country>
</aff>
<aff id="aff5">
<sup>5</sup>
<institution>Department of Forestry and Environmental Resources, North Carolina State University</institution>, <addr-line>Raleigh, NC</addr-line>, <country>United States</country>
</aff>
<aff id="aff6">
<sup>6</sup>
<institution>School of Environmental Science and Engineering, Tianjin University</institution>, <addr-line>Tianjin</addr-line>, <country>China</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited and Reviewed by: Huihui Li, Chinese Academy of Agricultural Sciences, China</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Juan Du, <email xlink:href="mailto:djuan@zju.edu.cn">djuan@zju.edu.cn</email>; Quanzi Li, <email xlink:href="mailto:20240082@zafu.edu.cn">20240082@zafu.edu.cn</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>04</day>
<month>12</month>
<year>2024</year>
</pub-date>
<pub-date pub-type="collection">
<year>2024</year>
</pub-date>
<volume>15</volume>
<elocation-id>1508507</elocation-id>
<history>
<date date-type="received">
<day>09</day>
<month>10</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>14</day>
<month>11</month>
<year>2024</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2024 Du, Ye, An, Chen, Wang, Wang, Lu and Li</copyright-statement>
<copyright-year>2024</copyright-year>
<copyright-holder>Du, Ye, An, Chen, Wang, Wang, Lu and Li</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="" journal-id-type="nlm-ta" xlink:href="https://www.frontiersin.org/research-topics/44714/research-advances-on-forest-tree-functional-genomics-and-breeding/overview" ext-link-type="uri">Editorial on the Research Topic <article-title>Research advances on forest tree functional genomics and breeding</article-title>
</related-article>
<kwd-group>
<kwd>forest tree</kwd>
<kwd>woody plants</kwd>
<kwd>functional genomics</kwd>
<kwd>breeding</kwd>
<kwd>genome editing</kwd>
</kwd-group>
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<fig-count count="0"/>
<table-count count="0"/>
<equation-count count="0"/>
<ref-count count="23"/>
<page-count count="3"/>
<word-count count="1195"/>
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<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Functional and Applied Plant Genomics</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<p>Forest ecosystems&#x2014;one of the biggest carbon sinks in the world&#x2014;play a key role in terrestrial biodiversity and carbon sequestration. As important sustainable resources, trees serve as rich sources of agronomic and economic traits, providing wood, pulp and paper, fiber-related products, energy, and chemical products. Over the past few decades, conventional crossbreeding has helped generate plant varieties with improved agronomic and economic traits. However, conventional crossbreeding in forestry is time-intensive and has reached a bottleneck. Thus, improvement in growth and agronomic and economically important traits in tree species requires attention. Biotechnology has recently resulted in great progress in crop breeding, owing to the development of high-quality genome assembly and annotation tools, gene identification techniques, and efficient gene editing. Nonetheless, compared to crop species, extensive efforts are needed for the assembly and annotation of high-quality genomes, identification of key genes regulating agronomic and economically important traits, and highly efficient gene editing in tree species that exhibit high heterozygosity.</p>
<p>This Frontiers Research Topic aims to present the latest fundamental discoveries in the field of forest tree genomics, including genetic studies focusing on the genes and pathways associated with key agronomic and economically important traits, molecular mechanisms underlying secondary growth regulation, and the potential utilization of biotechnology in genetic improvement of woody plant species.</p>
<p>This volume is organized into the following sections: (1) genome assembly and annotation; (2) functional identification of key genes regulating tree growth, vascular development, and stress response; and (3) genetic transformation and gene editing in woody plants.</p>
<sec id="s1">
<title>Genome assembly</title>
<p>The first tree species with a completely sequenced genome was <italic>Populus trichocarpa</italic> (<xref ref-type="bibr" rid="B20">Tuskan et&#xa0;al., 2006</xref>). With the increase in sequencing depth and length and decrease in sequencing cost, the genomes of several woody plants, including poplar, have been sequenced (<xref ref-type="bibr" rid="B13">Liu et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B14">Liu et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B7">Huang et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B17">Niu et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B21">Zhang et&#xa0;al., 2024</xref>). Researchers have also been able to sequence a nearly gapless, highly contiguous genome of a doubled haploid line of <italic>P. ussuriensis</italic>, whose 19 chromosomes have been assembled from telomere to telomere (<xref ref-type="bibr" rid="B12">Liu et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B16">Niu and Li, 2024</xref>; <xref ref-type="bibr" rid="B18">Shi et&#xa0;al., 2024</xref>). Additionally, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1218515">Sahu et&#xa0;al.</ext-link> have assembled the genome of Indian rosewood at the chromosome level, providing the foundation for studying tree growth and wood formation in this species. The assembly of this reference genome is likely to promote research on the formation of heartwood, which is an economically important wood.</p>
</sec>
<sec id="s2">
<title>Functional identification of key genes regulating the growth, vascular development, and stress response of tree species</title>
<p>Tree biomass is determined by longitudinal and lateral growth. The cultivation of fast-growing trees is of great interest to produce adequate biomass in a shorter time. In this regard, researchers have attempted to identify the key genes regulating the growth rate of tree species for genetic improvement. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1241908">Akutsu et&#xa0;al.</ext-link> used genomic selection (GS) for growth characteristics in an open-pollinated breeding population of Korean red pine (<italic>Pinus densiflora</italic>) and concluded that the trained GS model was more effective than traditional breeding methods. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2024.1285094">Kang et&#xa0;al.</ext-link> conducted GS prediction on a half-sib progeny population of <italic>Shorea macrophylla</italic> using six methods, and showed that GS with GWAS-based SNP selection was useful for breeding tree species.</p>
<p>Wood formation, or secondary growth, is a biological process specific to woody plants. The continuous activities of the cambium, comprising cambial cell proliferation, cell expansion, secondary wall thickening, and programmed cell death (PCD), produce xylem (wood) (<xref ref-type="bibr" rid="B4">Du et&#xa0;al., 2023</xref>). Key regulators of cambial cell proliferation and secondary wall thickening in poplar trees have been identified using forward and reverse genetics (<xref ref-type="bibr" rid="B6">Hu et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B9">Kim et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B15">Luo and Li, 2022</xref>; <xref ref-type="bibr" rid="B3">Dai et&#xa0;al., 2023</xref>; <xref ref-type="bibr" rid="B11">Li et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B21">Zhang et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B22">Zhou et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B23">Zhu and Li, 2024</xref>). However, studies on PCD in xylem cells are still limited.</p>
<p>
<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1196618">Liu et&#xa0;al.</ext-link> stained different xylem cells using different dyes and sorted living cells, early PCD cells, and late PCD cells using flow cytometry. This method can also improve the analysis of gene expression dynamics along the continuous developmental stages of PCD. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1302536">Gu&#xe9;rin et&#xa0;al.</ext-link> generated dominant repressor poplar lines for <italic>PtaPLATZ18</italic>, which encodes an A/T-rich and zinc-binding protein. <italic>PtaPLATZ18-SRDX</italic> transgenic lines exhibited wider xylem compared to the wildtype plants, with a higher lignin content in transgenic wood. Moreover, the transgenic plants exhibited significantly increased height, suggesting the potential of this gene in promoting tree growth and wood production.</p>
<p>Tree growth is immensely affected by environmental factors such as drought and high temperatures (<xref ref-type="bibr" rid="B8">Jiang et&#xa0;al., 2023</xref>), which induce several physiological and molecular processes, such as abscisic acid synthesis in roots and leaves. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1297235">Yu et&#xa0;al.</ext-link> analyzed the transcriptomic profile of <italic>Phoebe bournei</italic>, an important afforestation tree species in the subtropical region of China, exposed to drought stress. Through gene co-expression network analysis, they identified two core transcription factors, TGA4 and APRR2, involved in drought. Nonetheless, further genetic experiments are required to determine the functions of the candidate genes.</p>
</sec>
<sec id="s3">
<title>Genetic transformation and gene editing in woody plants</title>
<p>Genetic transformation is vital to characterize the function of a gene. In addition to genome assembly, successful genetic transformation using stems (<xref ref-type="bibr" rid="B19">Song et&#xa0;al., 2006</xref>) and leaves as explants (<xref ref-type="bibr" rid="B10">Li et&#xa0;al., 2017</xref>) makes <italic>P. trichocarpa</italic> a model species. As <italic>P. trichocarpa</italic> is an endemic species, researchers have resort to other poplar species, such as <italic>P. alba</italic>, <italic>P. alba</italic> &#xd7; <italic>P. glandulosa</italic>, and <italic>P. tremula</italic>. Transformation in poplar, which relies on tissue culture, is mostly mediated by <italic>Agrobacterium tumefaciens</italic>, although <italic>A. rhizogenes</italic>-mediated transformation is independent of tissue culture and produces hairy roots, from which plants can regenerate. In this regard, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1196561">Ying et&#xa0;al.</ext-link> reviewed recent advances in <italic>A. rhizogenes</italic>-mediated transformation and Ri breeding in woody plants and emphasized its potential application in the difficult-to-propagate woody species.</p>
<p>Gene editing has been increasingly applied for the genetic improvement of plant species because of its high precision (<xref ref-type="bibr" rid="B2">Borthakur et&#xa0;al., 2022</xref>). Fan et&#xa0;al. first used the type II clustered regularly interspaced short palindromic repeats (CRISPR)-associated protein (Cas9) system in <italic>P. tomentosa</italic> (<xref ref-type="bibr" rid="B5">Fan et&#xa0;al., 2015</xref>). Furthermore, the CRISPR-Cas12 system has been used to knock out multiple targets of <italic>Phytoene desaturase 8</italic> in poplar (<xref ref-type="bibr" rid="B1">An et&#xa0;al., 2020</xref>). In this study, An et&#xa0;al. also evaluated the effects of temperature on gene editing efficiency and observed that the majority of editing included large-fragment deletions. Similarly, <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fpls.2023.1159615">Movahedi et&#xa0;al.</ext-link> examined the effects of three factors on editing and showed that high <italic>Agrobacteria</italic> concentration, increased DDT number, and optimized homologous arm length resulted in efficient homology-directed repair. Currently, the CRISPR-Cas system is most commonly used for gene editing in poplar, but different woody plant species may need different CRISPR-Cas systems. Thus, increased efforts are needed to test the efficiency of gene editing in woody plants other than poplar.</p>
<p>This Research Topic incorporates articles on genome assembly and annotation; functional identification of key genes regulating tree growth, vascular development, and stress response; and genetic transformation and gene editing in woody plants, and serves as a valuable resource for studying forest tree breeding and functional genomics.</p>
</sec>
</body>
<back>
<sec id="s4" sec-type="author-contributions">
<title>Author contributions</title>
<p>JD: Writing &#x2013; review &amp; editing, Writing &#x2013; original draft. YA: Validation, Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, Supervision, Visualization, Writing &#x2013; review &amp; editing. YC: Validation, Writing &#x2013; review &amp; editing. JPW: Writing &#x2013; review &amp; editing. JW: Validation, Writing &#x2013; review &amp; editing. ML: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization, Writing &#x2013; review &amp; editing. QL: Writing &#x2013; original draft, Writing &#x2013; review &amp; editing. TY: Data curation, Writing &#x2013; review &amp; editing.</p>
</sec>
<sec id="s5" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare financial support was received for the research, authorship, and/or publication of this article. This work is supported by the National Natural Science Foundation of China (32071792), National Key Program on 2023YFE124800, and the Key program of the National Science Foundation of Zhejiang province (LZ22C160002) to JD, Zhejiang University.</p>
</sec>
<sec id="s6" 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="s7" 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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