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Cytogenetics, Genome Size and Anther Anatomy in Bulbostylis (Cyperaceae)

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Cytogenetics, Genome Size and Anther Anatomy in Bulbostylis (Cyperaceae) Caryologia International Journal of Cytology, Cytosystematics and Cytogenetics ISSN: 0008-7114 (Print) 2165-5391 (Online) Journal homepage: http://www.tandfonline.com/loi/tcar20 Cytogenetics, genome size and anther anatomy in Bulbostylis (Cyperaceae) Maria Gabriela López, Fernández Aveliano & Galdeano Florencia To cite this article: Maria Gabriela López, Fernández Aveliano & Galdeano Florencia (2017): Cytogenetics, genome size and anther anatomy in Bulbostylis (Cyperaceae), Caryologia, DOI: 10.1080/00087114.2017.1335515 To link to this article: http://dx.doi.org/10.1080/00087114.2017.1335515 Published online: 28 Jun 2017. Submit your article to this journal 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: [181.15.152.229] Date: 28 June 2017, At: 12:03 CARYOLOGIA: INTERNATIONAL JOURNAL OF CYTOLOGY, CYTOSYSTEMATICS AND CYTOGENETICS, 2017 https://doi.org/10.1080/00087114.2017.1335515 Cytogenetics, genome size and anther anatomy in Bulbostylis (Cyperaceae) Maria Gabriela Lópeza,b, Fernández Avelianob and Galdeano Florenciaa,b aFacultad de Ciencias Agrarias, FCA-UNNE, Corrientes, Argentina; bInstituto de Botánica del Nordeste, UNNE-CONICET, Corrientes, Argentina ABSTRACT ARTICLE HISTORY Cyperaceae display diverse and unusual cytogenetical characteristics, but are still sparsely Received 8 February 2017 documented and poorly understood in this respect. Six new chromosome counts are presented Accepted 22 May 2017 for Bulbostylis. Absolute DNA content was close to 1 pg in all species and was independent KEYWORDS from chromosome number. Despite variations in chromosome number from 2n = 30 to Bulbostylis; holokinetic 2n = 102, DNA content only varied from 0.932 pg to 1.198 pg. The studies presented in this chromosomes; work showed that species possess holokinetic chromosomes. The size of stomata and pollen agmatoploidy; symploidy; grains did not vary. These results would confirm that the chromosome evolution of these species pseudomonads was via agmatoploidy and symploidy rather than by polyploidy. Abnormalities during meiotic spindle formation in microsporogenesis followed by simultaneous cytokinesis resulted in four microspores, of which only one was functional, resulting in a pseudomonad maturing into pollen grains with the usual vegetative and generative nuclei in addition to three degenerated microspore nuclei. Introduction to observe the remaining chiasmata. Because the sep- Family Cyperaceae has a cosmopolitan distribution with aration occurs between sister chromatids, the division a significant concentration of genera in the tropics. It is equational (Nordenskiöld 1962). In contrast, in the includes about 5500 species in 109 genera (Govaerts anaphase I of monocentric chromosomes the separation is between homologous chromosomes. In Cyperaceae, et al. 2007), 14 tribes and four subfamilies (Goetghebeur 1998). Most species occur in wet habitats or submerged holokinetic behavior has been studied only in Carex fields. Cyperaceae is a well-defined monophyletic (Wahl 1940) and Eleocharis palustris (Hakansson 1954). group (Muasya et al. 1998, 2000) included in Poales In the sister family Juncaceae, it is known only from (Angiosperm Phylogeny Group A. P. G. III, 2009) Luzula purpurea (Nordenskiöld 1962). and sister to Juncaceae as supported by morphologi- The absence of a single primary constriction and cal and molecular studies (Simpson 1995; Bruhl 1995; thus of a localized centromere has been termed in var- Goetghebeur 1998; Muasya et al. 1998, 2000, 2009). ious ways, including “holocentric chromosomes”, “with Cyperaceae display peculiar cytogenetic characteris- diffuse centromere or kinetochore” (Schrader 1935), or tics. They have holokinetic chromosomes (i.e. the kinetic “non-localized centromere”. However, the term “holo- activity extends along the entire chromosome), and pol- centric chromosomes” is the most appropriate (Bauer len grains that develop from pseudomonads (Wahl 1940; 1952; Sybenga 1972), because it implies that the kinetic Malheiros-Gardé and Gardé 1950, 1951; Hakansson activity is distributed along the entire length of the chro- 1954; Nordenskiöld 1962). Holokinetic chromosomes mosome (=holokinetic) and not limited to a discrete can be recognized by the variability of chromosomic region, as it is in monocentric (=monokinetic) chromo- fragments and the presence of a large kinetochore plate. somes (Papeschi 1992). Therefore, the term “holokinetic Schrader (1935) was the first to observe the peculiar chromosomes” reflects both the morphology and behav- movement of chromosomes in the mitotic anaphase and ior of chromosomes (Mola and Papeschi 2006). to use the term “diffuse centromere”. Karyotypic evolution in Cyperaceae is not only by Many organisms with holokinetic chromosomes have polyploidy, but also by agmatoploidy (i.e. fragmentation an uncommon meiotic behavior. The behavior is related of chromosomes) and symploidy (i.e. fusion of chro- to the orientation and migration of chromatids, which is mosomes). The term agmatoploidy was first used in the characteristic of holokinetic chromosomes. The differ- genus Luzula by Malheiros-Gardé and Gardé (1950). ence is observed at anaphase I: chromatids individualize, The occurrence of fragmentations (agmatoploidy) as if they were distinct units, and sister chromatids move and fusions (symploidy) of chromosomes in Cyperaceae apart from each other. In this phase, it is still possible makes it difficult to determine the basic chromosome CONTACT Maria Gabriela López [email protected] © 2017 Dipartimento di Biologia Evoluzionistica, Università di Firenze .L. MARIA GABRIELA ET AL 2 number. However, the existence of several genera and Capital, Laguna Soto, lomada arenosa. 16 July 1991. M. species with chromosome numbers that are multiples G. López 57 (CTES). of five suggests that the ancestral basic number in the B. hirtella (Schrad.) Nees ex Urb. Argentina, family is x=5, while other basic numbers, such as x=2, Corrientes, Dpto. Monte Caseros, desembocadura del 4, 6, and 12 are possible (Luceño et al. 1998a). río Miriñay y río Uruguay. Barranca arenosa. 12 October Another poorly studied aspect of Cyperaceae cytol- 2008. López 391 (CTES). ogy is DNA content. DNA content has been measured B. juncoides (Vahl) Kük. ex Herter var. juncoides. only in a few species (Bennet and Leitch 1995): Cyperus Argentina, Córdoba, Dpto. San Alberto, Pampa de Achala; haspan, C. textiles, Eleocharis acicularis, Eriophorum borde de Arroyo. 14 January 2009. López 388 (CTES). vaginatum, Schoenoplectus lacustris (=Scirpus lacus- B. rugosa M. G. López. Argentina, Corrientes, Dpto. tris) and Schoenoplectus tabernaemontani (=Scirpus Empedrado, frente al camping Municipal; barranca tabernaemontani). arenosa erosionada. 12 October 2009. López 389 (CTES). In this study, we focus on the genus Bulbostylis of tribe B. sphaerocephala (Boeckeler) Lindm. var. sphaero- Abildgaardieae. Kral (1971) focused on the cytology at cephala. Argentina, Misiones, Dpto. San Ignacio, Teyú the tribe level and is the only published study includ- Cuaré. 12 February 2009. López 394 (CTES). ing American Bulbostylis. Since he found that the basic B. sphaerolepis (Boeckeler) Beetle. Argentina, number is x = 5, he concludes that polyploidy represents Corrientes, Dpto. Empedrado, frente al camping the evolutionary mechanism explaining the cytological Municipal; barranca. 14 March 2006. López 378 (CTES). diversity of the tribe and considers Fimbristylis spathacea B. subtilis M. G. López. Argentina, Misiones, Dpto. an exception derived from another basic number, x = 6, 8. Eldorado, ruta provincial 17, paraje Cerro 60, cantera Despite his study, cytogenetic studies in Bulbostylis remain abandonada; entre piedras. 12 March 2009. López 393 scarce (see Roalson 2008), and chromosome counts are (CTES). known from only 16 of its 150 species (10.6% of the genus; see literature survey in Table 1). To fill this gap, we studied Methods the cytology, microsporogenesis, and microgametogene- sis of seven southern South American Bulbostylis species. The ploidy level of the seven studied species (see names In particular, we determined the mitotic chromosome and accession information in material studied) was number, the genome size, pollen grain and stomata size determined by counting chromosomes (eight cells per (to identify any correlation with chromosome numbers), species) in meristematic cells of root tips. Roots were and estimated pollen viability. Finally, we also described pretreated in a saturated solution of α-bromonaphtha- anther wall and pollen formation. lene for 2 h, hydrolyzed in HCl (1 N) for 10 min at 60 °C, and stained with Schiff’s reagent. Finally, the root Material and methods tips were smashed on a glass slide in a drop of 2% acetic orcein and analyzed with light microscopy. Material studied For measurements of genome size, small leaf samples Bulbostylis communis M.G. López & D. Simpson var. of similar size were taken from Bulbostylis plants and communis Argentina, Corrientes, Dpto. Monte Caseros, the standard Paspalum dilatatum Poir. ssp. flavescens desembocadura del río Miriñay y río Uruguay; Playa Roseng. (Q 3952), and then placed together in a Petri arenosa. 12 March 2009. López 398 (CTES). Dpto. dish (96 mm) with 0.5 ml nuclear extraction buffer from Table 1. Chromosome numbers in Bulbostylis from previously published studies. Species 2n Source Bulbostylis barbata (Rottb.) C.B. Clarke 10
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