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Karyotype Characterization of Four Mexican Species of Schoenoplectus (Cyperaceae) and First Report of Polyploid Mixoploidy for the Family

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Karyotype Characterization of Four Mexican Species of Schoenoplectus (Cyperaceae) and First Report of Polyploid Mixoploidy for the Family Caryologia International Journal of Cytology, Cytosystematics and Cytogenetics ISSN: 0008-7114 (Print) 2165-5391 (Online) Journal homepage: http://www.tandfonline.com/loi/tcar20 Karyotype characterization of four Mexican species of Schoenoplectus (Cyperaceae) and first report of polyploid mixoploidy for the family Jorge A. Tena-Flores, M. Socorro González-Elizondo, Yolanda Herrera-Arrieta, Norma Almaraz-Abarca, Netzahualcóyotl Mayek-Pérez & André L.L. Vanzela To cite this article: Jorge A. Tena-Flores, M. Socorro González-Elizondo, Yolanda Herrera- Arrieta, Norma Almaraz-Abarca, Netzahualcóyotl Mayek-Pérez & André L.L. Vanzela (2014) Karyotype characterization of four Mexican species of Schoenoplectus (Cyperaceae) and first report of polyploid mixoploidy for the family, Caryologia, 67:2, 124-134, DOI: 10.1080/00087114.2014.931633 To link to this article: http://dx.doi.org/10.1080/00087114.2014.931633 Published online: 20 Aug 2014. Submit your article to this journal Article views: 51 View related articles View Crossmark data Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=tcar20 Download by: [148.204.124.71] Date: 23 February 2017, At: 09:58 Caryologia: International Journal of Cytology, Cytosystematics and Cytogenetics, 2014 Vol. 67, No. 2, 124–134, http://dx.doi.org/10.1080/00087114.2014.931633 Karyotype characterization of four Mexican species of Schoenoplectus (Cyperaceae) and first report of polyploid mixoploidy for the family Jorge A. Tena-Floresa*, M. Socorro González-Elizondoa, Yolanda Herrera-Arrietaa, Norma Almaraz-Abarcaa, Netzahualcóyotl Mayek-Pérezb and André L.L. Vanzelac aCIIDIR Durango, Instituto Politécnico Nacional, Durango, Durango, Mexico; bCentro de Biotecnología Genómica, Instituto Politécnico Nacional, Reynosa, Tamaulipas, México; cLaboratório de Citogenética e Diversidade Vegetal, Departamento de Biologia Geral, CCB, Universidade Estadual de Londrina, Londrina, Paraná, Brazil Karyomorphological analysis of four species of Schoenoplectus (Cyperaceae) from north-central Mexico were carried out. Chromosome numbers ranged from 2n =38to2n = 84. New records of counting are given for Schoenoplectus acutus var. occidentalis (2n = 38 and 2n = 84) and S. americanus (2n = 66). Intra-individual variation in chromosome number is reported for the first time for S. acutus, with a rare polyploid mixoploidy with a prevalence of cells with 2n = 38 (36 small + 2 compound, larger chromosomes) and a few cells with 2n = 84 small, dot-shaped chromosomes, this being the first record of polyploid mixoploidy for Cyperaceae. Mean length of the diploid set ranged from 51.5 μm (S. tabernaemontani) to 79.5 μm(S. acutus). The lowest average chromosome length for the dot-shaped chromosomes was 0.69 μm(S. acutus) and the highest 1.62 μm(S. tabernaemontani); the pair of large chromosomes in S. acutus reached 3.17 μm. A low interchromosomal asymmetry index (A2), 0.11 to 0.14 was found, very similar among all the species except for S. acutus (A2 = 0.30). Absence of primary constrictions was confirmed. The most common mecha- nism of karyotype variation in the studied species is dysploidy, followed by polyploidy. A comparison of chromosome numbers between Schoenoplectus and the recently segregated Schoenoplectiella based on the literature reveals that Schoenoplectus has higher numbers (n = 18 to 64; 2n = 36 to 84) than Schoenoplectiella (n = 5 to 44; 2n = 18 to 76) as well as a higher prevalence of disploids. Keywords: dysploidy; holocentric chromosomes; mixoploidy; polyploidy; Schoenoplectiella; Scirpus; sedges Introduction aquatic vegetation. They have a great environmental Schoenoplectus (Rchb.) Palla is a genus of Cyperaceae value (for phytoremediation, as soil stabilizers, habitat distributed almost worldwide, with about 31 species and and food for wild species), as well as important ethnobo- five nothospecies, not including those that are at present tanic and economic values, e.g. for S. californicus (C. A. considered under Bolboschoenus (Asch.) Palla and Meyer) Soják, a species widely distributed in the New Schoenoplectiella Lye (Lye 2003). Schuyler noted since World (Mexican tules, Peruan and Bolivian totora). In 1976 that “the genus Scirpus (sensu lato) is a diverse North America their culms are used, mostly historically, assemblage of unrelated species groups which eventually for making mats, baskets, chair seats, houses, boats, and will be treated as different genera”. The three above- other objects. Some species are cultivated as ornamentals mentioned genera have been segregated from Scirpus L. (Smith 2002). Five species of Schoenoplectus s.s. are and are provisionally considered in the tribe Fuireneae known in Mexico, where some of them are used for craft Reichenb. ex Fenzl (Goetghebeur 1998; Muasya, making and phytoremediation (González et al. 2007). Vrijdaghs, et al. 2009), although further studies are in Except for two Japanese studies (Maeda and Uchino progress to evaluate their relationships (Muasya, 2004; Yano and Hoshino 2005) very few cytogenetic Vrijdaghs, et al. 2009). studies have been performed recently on Schoenoplectus, fi Groups in Schoenoplectus and related genera have and the rst of them was focused on species now recog- been distinguished based on morphology and micromor- nized under Schoenoplectiella. phology (Strong 1993, 1994; Bruhl 1995; Goetghebeur Like the other members of Cyperaceae, Schoenoplec- “ ” 1998; Pignotti and Mariotti 2004), as well as molecular tus has holokinetic ( holocentric ) chromosomes. data (Muasya et al. 2000; Young et al. 2002; Yano and Although they are sometimes thought to be rare, holoki- Hoshino 2005; Jung and Choi 2010), revealing the group netic chromosomes are rather common among many to be paraphyletic or polyphyletic. Schoenoplectus groups of organisms, including two plant families: includes some difficult species complexes and is in need Cyperaceae and Juncaceae (Luceño et al. 1998). Diffuse of a systematic revision (Smith 2002; Shiels and Monfils kinetochores favor karyotype differentiation via agmatop- fi 2012). It is frequently found in wetlands and marshes, loidy ( ssion), symploidy (fusion), and polyploidy and some species are often the dominant element in (Luceño and Guerra 1996) as well as chromosome *Corresponding author. Email: jorge@tena-flores.com © 2014 Dipartimento di Biologia Evoluzionistica, Università di Firenze Caryologia: International Journal of Cytology, Cytosystematics and Cytogenetics 125 viability after most rearrangements (Guerra 2008). For h, and stored at –20°C until use. Root tips were washed this reason, chromosomes evolve very dynamically in in distilled water, digested for 1 h in a mixture of 4% Cyperaceae (Luceño and Guerra 1996; Hipp et al. 2009) (w:v) cellulase and 40% (w:v) pectinase at 37°C, further and in wood-rushes (Luzula DC., Juncaceae) (Bozek hydrolyzed in 1 M HCl for 10 min at 60°C, washed et al. 2012), where a considerable diversity of karyotype again in distilled water, and squashed in a drop of 45% evolution occurs. acetic acid. The cover slips were removed after freezing Given that holokinetic chromosomes lack a defined in liquid nitrogen. Slides were stained in 4% hematoxy- centromere they have no primary constriction, so the lin and permanent slides mounted in Entellan (Merck options for analysis based on morphology of their karyo- KGaA, Darmstadt, Germany). Chromosome counts were types are greatly reduced. Karyotype parameters, such as made in at least 20 cells. Mean lengths of the karyotype intrachromosomal asymmetry, can only be estimated (the total diploid length) and of the shortest and longest using the chromosome length; however, the high varia- chromosome of the complement were calculated. Chro- tion in chromosome number, interchromosomal asymme- mosome measurements were carried out using the free- try index, and presence or absence of nucleolar ware computer application MicroMeasure software constrictions are useful for karyotype description. version 3.3 for Windows (Colorado State University, Mexican Cyperaceae are poorly known cytologically. Dept of Biology, USA)(http://www.colostate.edu/Depts/ Here, we present karyomorphological data of four species Biology/MicroMeasure) and the data were used to of Schoenoplectus (Cyperaceae) from north-central Mexico. assemble ideograms. For each sample, from 5 to 10 metaphase spreads with similar condensation were Materials and methods measured. All images were acquired with a Carl Zeiss AxioImag- Mitotic metaphase chromosomes were studied from root er.Z2 microscope (ZEISS. Oberkochen, Germany) meristematic cells. Eleven populations representing four equipped with an Axiocam Hrc camera and AxioVs40 species of Schoenoplectus were collected in seven locali- Rel.4.8.2 software (ZEISS. Oberkochen, Germany). Inter- ties of north-central Mexico, in the state of Durango. chromosomal asymmetry was calculated using the Romer- Voucher specimens were deposited in herbarium CIIDIR o-Zarco (1986) index based on Pearson’s dispersion at the Centro Interdisciplinario de Investigación para el coefficient (the ratio between the standard deviation (s) Desarrollo Integral Regional, Instituto Politécnico Nac- and the mean of chromosome length ðXÞ for each sample): ional, in Durango, Mexico. Samples were cultivated in the greenhouse of CIIDIR. Data of the studied taxa, ¼ s A2 (1) localities of collection and voucher specimens are pre- X sented in Table 1. The karyotypical analyses were performed with root Results tips pre-treated
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