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Phylogeny and Maternal Donors of Elytrigia Desv. Sensu Lato (Triticeae

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Phylogeny and Maternal Donors of Elytrigia Desv. Sensu Lato (Triticeae Yang et al. BMC Plant Biology (2017) 17:207 DOI 10.1186/s12870-017-1163-7 RESEARCH Open Access Phylogeny and maternal donors of Elytrigia Desv. sensu lato (Triticeae; Poaceae) inferred from nuclear internal-transcribed spacer and trnL-F sequences Yan Yang1,2, Xing Fan1, Long Wang1, Hai-Qin Zhang1, Li-Na Sha1, Yi Wang1, Hou-Yang Kang1, Jian Zeng3, Xiao-Fang Yu4 and Yong-Hong Zhou1* Abstract Background: Elytrigia Desv. is a genus with a varied array of morphology, cytology, ecology, and distribution in Triticeae. Classification and systematic position of Elytrigia remain controversial. We used nuclear internal-transcribed spacer (nrITS) sequences and chloroplast trnL-F region to study the relationships of phylogenetic and maternal genome donor of Elytrigia Desv. sensu lato. Results: (1) E, F, P, St, and W genomes bear close relationship with one another and are distant from H and Ns genomes. Ee and Eb are homoeologous. (2) In ESt genome species, E genome is the origin of diploid Elytrigia species with E genome, St genome is the origin of Pseudoroegneria. (3) Diploid species Et. elongata were differentiated. (4) Et. stipifolia and Et. varnensis sequences are diverse based on nrITS data. (5) Et. lolioides contains St and H genomes and belongs to Elymus s. l. (6) E genome diploid species in Elytrigia serve as maternal donors of E genome for Et. nodosa (PI547344), Et. farcta, Et. pontica, Et. pycnantha, Et. scirpea,andEt. scythica. At least two species act as maternal donor of allopolyploids (ESt and EStP genomes). Conclusions: Our results suggested that Elytrigia s. l. species contain different genomes, which should be divided into different genera. However, the genomes of Elytrigia species had close relationships with one another. Diploid species were differentiated, because of introgression and different geographical sources. The results also suggested that the same species and the same genomes of different species have different maternal donor. Further study of molecular biology and cytology could facilitate the evaluation of our results of phylogenetic in a more specific and accurate way. Keywords: Elytrigia Desv., Chloroplast trnL-F, Nuclear ITS, Phylogeny, Maternal donor Background As one of the most important perennial genera of Triticeae in Poaceae includes not only the most econom- Triticeae, Elytrigia Desv. includes 40 species, which are ically important cereal crops (wheat, barley, and rye) but distributed in subtropical and warm temperate regions also forage grasses and ecological species in grasslands. of both hemispheres [6]. Elytrigia Desv. was established Approximately 450 Triticeae species exist worldwide [1– by Desvaux [7], with Elytrigia repens (L.) Nevski as the 3]. Given the wide variety of biological mechanisms and type species. Morphologically, Elytrigia sensu lato is genetic systems, this tribe represents an excellent group characterized by branched creeping rhizomes and for research on plant systematics, genetic diversity, and caespitose, long anthers, lanceolate to liner glumes, speciation [4, 5]. lanceolate lemma, single spikelet per node, and cross- pollination, and most species were previously categorized under Agropyron Gaertner [1, 6, 8]. * Correspondence: [email protected] 1Triticeae Research Institute, Sichuan Agricultural University, Wenjiang, Cytogenetically, ploidy levels in Elytrigia s. l. vary from 611130 Chengdu, Sichuan, People’s Republic of China diploid (2n = 2× = 14) to decaploid (2n = 10× = 70) and Full list of author information is available at the end of the article © The Author(s). 2017 Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. Yang et al. BMC Plant Biology (2017) 17:207 Page 2 of 14 contain Ee,Eb, St, ESt, StH, and NsXmStH genomes [1, genomes [28, 39]. nrITS sequences were widely applied 3, 8–11]. According to the proposed genomic system of to explain genomic and phylogenetic relationship at a classification, Löve [8] suggested that Elytrigia s. l. low taxonomic levels [40–42] and Triticeae species con- approximately includes 60 species and varieties and di- taining E, H, Ns, P, St and Xm genomes in Elymus, vided them into five genera, namely, Pseudoroegneria Hordeum, Psathyrostachys, Agropyron Pseudoroegneria (Nevski) Á. Löve (St), Lophopyrum Á. Löve (E), Thino- and Leymus [5, 27, 33, 40–43]. Chloroplast DNA pyrum Á. Löve (J), Elytrigia (EJSt), and Elymus (StH). (cpDNA) sequences, including intron of trn-L and inter- Dewey [1] considered Elytrigia s. l. into three independ- genic spacer of trnH-psbA, trnL-trnF, and trnS-trnG, are ent genera: Pseudoroegneria (St), Thinopyrum (E or J), also widely used to identify maternal donors of poly- and Elytrigia (StX). Studies showed similarity of the E ploids with extra ability to analyze phylogenetic relation- and J genomes [12–19]. Wang et al. [19] suggested that ships among relevant species [41, 44–47]. E and J should be considered as identical genomes and The present study analyzed sequence data of one ITS be distinguished from Ee and Eb. With the genomic sys- region of nuclear DNA and one chloroplast gene (the tem of classification in Triticeae taxonomy and system- intergenic region of trnL-trnF) from 18 species (subspe- atics and genomic constitutions of increasing species cies) in Elytrigia s. l. with 21 species of related genera in identified, the definition of Elytrigia becomes narrower Triticeae. The objectives are as follows: (1) to investigate than that of traditional Elytrigia s. l. and only includes phylogenetic relationships among species in Elytrigia s. l. all polyploidy taxa with combination of Ee,Eb, and St ge- and related genera; (2) to elucidate interspecific nomes [3, 9, 11]. Ee genome originated from Lopho- relationships among Elytrigia s. l. species; (3) to study pyrum elongatum (Host) Á. Löve, Eb genome from phylogenetic relationships among species with different Thinoopyrum bessarabicum (Savul. and Rayss) Á. Löve, genomes and genome combinations; and (4) to discuss and St genome from diploid species in Pseudoroegneria putative maternal donors for ESt genome species. (Nevski) Á. Löve. However, some studies reported that Et. repens, a type of Elytrigia, is a hexaploid species with Methods StStStStHH genomes and was renamed as Elymus repens Plant materials [3, 11, 20, 21]. Therefore, definition, precise taxonomic This study included 18 species (subspecies) of Elytrigia ranks, and number of Elytrigia species remain Desv. sensu lato and 21 species (subspecies) of related controversial. genera in Triticeae. Table 1 lists names, accession num- Polyploidization and hybridization are the two main bers, genomes, origins, and GenBank accession num- mechanisms in plant speciation and evolution [22, 23]. bers. Bromus cartharticus Vahl. and Bromus tectorum L. The changes in the cell size, genome size, gene expres- were used as outgroup [5, 41, 48]. Seed materials with sion, genomic repatterning, epigenetic effects and retro- W6 and PI accession numbers were carefully offered by transposon activation are caused by the polyploidization the American National Plant Germplasm System and chromosome doubling [5, 22–26]. As a result of (Pullman, Washington, USA). We gathered seed mate- these changes, stabilization of hybrid condition and full rials with Y and ZY numbers. Voucher specimens and fertility may occur. And the establishment of phenotypes plants were deposited at the perennial nursery and herb- in nature could be enhanced. Therefore, polyploids arium of the Triticeae Research Institute, Sichuan could adjust to match with the new ecological niches or Agricultural University, China. become more competitive than parental donors [5, 23, 26, 27]. The evolution of polyploidization alone and/or DNA extraction, amplification, and sequencing the combined effects of hybridization and polyploidiza- Total genomic DNA was extracted from leaves of single tion may lead to complex lineages, requiring an explan- plants by slight modification of Cetyltrimethyl Ammo- ation of the phylogenetic relationship [27]. Molecular nium Bromide (CTAB) procedure [49]. nrITS sequence genetic analysis bears significance in elucidating phylo- and chloroplast trnL-F sequence were amplified with genetic relationships and genome evolution patterns in primers described in Table 2. A final volume of 20 μLof taxa for these kinds of plant groups [27, 28]. The ana- mixed reagents was obtained for each polymerase chain lysis of Molecular phylogenetic exploits DNA sequences reaction (PCR); reagents included 2× Taq PCR Master- elucidated the history of revolution and origins of spe- Mix (10× ExTaq polymerase buffer, 3 mmol/L MgCl2, cies in Triticeae. This illustrates their hybridization 500 μmol/L deoxynucleotide, 100 mmol/L KCl, and events and parental lineages contains their formation, 20 mmol/L Tris–HCl), 1 μmol/L of each primer, and identifies the polyploidization mode [29–38]. Repro- 20–40 ng of template DNA, and distilled deionized ducibility and simplicity represent the main qualities that water. PCR reactions were performed
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