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First Cytogenetic Register of an Allopolyploid Lineage of the Genus Aeschynomene Citation: F

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First Cytogenetic Register of an Allopolyploid Lineage of the Genus Aeschynomene Citation: F Firenze University Press Caryologia www.fupress.com/caryologia International Journal of Cytology, Cytosystematics and Cytogenetics First cytogenetic register of an allopolyploid lineage of the genus Aeschynomene Citation: F. Tapia-Pastrana, A. Delga- do-Salinas (2020) First cytogenetic register (Leguminosae, Papilionoideae) native to Mexico of an allopolyploid lineage of the genus Aeschynomene (Leguminosae, Papil- ionoideae) native to Mexico. Caryolo- gia 73(4): 17-26. doi: 10.13128/caryolo- Fernando Tapia-Pastrana1,*, Alfonso Delgado-Salinas2 gia-949 1 Facultad de Estudios Superiores Zaragoza, Universidad Nacional Autónoma de México, Received: May 24, 2020 Laboratorio de Genecología, Batalla 5 de Mayo s/n esquina Fuerte de Loreto, Col. Ejér- cito de Oriente, Iztapalapa, C.P. 09230, Ciudad de México, Mexico Accepted: November 10, 2020 2 Instituto de Biología, Departamento de Botánica, Universidad Nacional Autónoma de Published: May 19, 2021 México, Apartado Postal 70-233, 04510, Cd. de México, Mexico *Corresponding author. E-mail: [email protected] Copyright: © 2020 F. Tapia-Pastrana, A. Delgado-Salinas. This is an open access, peer-reviewed article pub- Abstract. A conventional cytogenetics analysis revealed for first time an allopolyploid lished by Firenze University Press lineage of the genus Aeschynomene in Mexico. The hybrid condition is confirmed after (http://www.fupress.com/caryologia) all the prometaphase and metaphase nuclei of the hybrids exhibited only one pair of and distributed under the terms of the SAT-chromosomes, confirming the existence of nucleolar dominance and amphiplasty. Creative Commons Attribution License, The karyotype formula for this lineage was 2n = 4x = 40 = 34 m + 6 sm with a total which permits unrestricted use, distri- diploid chromosome length (TDCL) = 28µm and an average chromosome size (AC) = bution, and reproduction in any medi- 1.40 µm. Comparison of the karyotype and other chromosomal parameters with recent um, provided the original author and source are credited. cytogenetics records for other species of the subgenus Aeschynomene included in the Nod-independent clade allows propose to Aeschynomene evenia and A. scabra as possi- Data Availability Statement: All rel- ble progenitors. Furthermore, other comparison of seedlings focused at the number of evant data are within the paper and its leaflets of the first four eophylls of the proposed parents and of the hybrid individuals Supporting Information files. allowed to observe coincidences that support the proposal made from the cytogenetic analysis. Evidence of “gigas” effects on flowers and fruits of hybrids is also shown. Competing Interests: The Author(s) declare(s) no conflict of interest. Keywords: cryptic taxa, cytotype, karyotype, nucleolar dominance, SAT-chromo- ORCID somes, secondary constrictions, seedlings. FTP: 0000-0003-0232-2110 I. INTRODUCTION Aeschynomene Linnaeus (Leguminosae, Tribe Dalbergieae s. l.) is a diverse genus of subfamily Papilionoideae (Papilionoid legumes) distributed in the tropics and subtropics of the world (Lavin et al. 2001, Klitgaard and Lavin 2005). It comprises herbaceous and woody species, annual, repetitive and perennial with different ecological requirements. Several species con- tribute to supplement nitrogen to the soil through the production of nodular roots and stems in symbiosis with nitrogen fixing bacteria, so they are eco- nomically important as green manure (Alazar and Becker 1987; Fernandes 1996; Souza et al. 2012) and recently, Aeschynomene evenia C. Wright has been proposed as a model species in genetics to develop new agronomic Caryologia. International Journal of Cytology, Cytosystematics and Cytogenetics 73(4): 17-26, 2020 ISSN 0008-7114 (print) | ISSN 2165-5391 (online) | DOI: 10.13128/caryologia-949 18 Fernando Tapia-Pastrana, Alfonso Delgado-Salinas strategies in the engineering of nitrogen fixing nodules J. Léonard (2n = 8x = 76) and A. schimperi Hochst. ex that enhance rice production (Arrighi et al. 2012, 2013). A.Rich. (2n = 8x = 56), where variations in the number This taxon belongs to the group of 11 semi-aquatic spe- of chromosomes also indicated possible dysploidy/ane- cies of Aeschynomene that have the property of being uploidy events. In Mexico, Aeschynomene is represented nodulated by photosynthetic Bradyrhizobium that lack by 31 species and infraspecific taxa including several the nodABC genes necessary for the synthesis of Nod endemisms. An investigation about the patterns of chro- factors and are grouped into the so-called Nod-inde- mosomal evolution in Mexican species, including six pendent clade (Chaintreuil et al. 2013; Brottier et al. taxa of the Nod-independent clade, showed the predomi- 2018) and that correspond to the morphological series nant of a basic 2n = 20 diploid structure and evolution- Indicae and Sensitivae (Rudd 1955). ary patterns related to the corresponding morphological The genus Aeschynomene traditionally included in series (Tapia-Pastrana et al. 2020). the Aeschynomeneae tribe (Polhill et al.1981) and cur- In the present research, a conventional cytogenetic rently circumscribed in the Dalbergioid clade (Lavin et study was carried out to obtain the karyotype and ana- al. 2001; Wojciechowski et al. 2004) has evolved in dif- lyze the level of ploidy in a Mexican population initially ferent ecological niches and includes herbaceous forms, described as Aeschynomene scabra G. Don, where the annual and perennial shrubs and trees up to 8 meters, size of the flowers, fruits and seeds generated suspicions with compound pinnate leaves and papilionoid flowers about a possible hybrid origin. In addition as the sam- that are generally self-pollinated, although there is cross- pled individuals exhibited floral morphotypes similar to pollination by bees (Rudd 1955; Fernandes 1996; Arrighi those of A. evenia C. Wright and A. scabra, whose col- et al. 2014, Carleial et al. 2015). Other studies indicate lection records in Mexico would support their partici- that the genus Aeschynomene is not monophyletic and pation in the hybridization process, the growth pattern taxa with basifixed stipules and a campanulate calyx of the first four eophylls was also compared in putative (subgenus Ochopodium Vogel) are more related to the hybrids and their parental assumptions. genera Machaerium Persoon and Dalbergia Linnaeus f. than to taxa with medfixed stipules and a bilabiate calyx (subgenus Aeschynomene Léonard) (Ribeiro et al. 2007; 2. MATERIAL AND METHODS Cardoso et al. 2012). Currently Aeschynomene genus contains 170 (http:// 2.1 Collection sites www.theplantlist.org) to 180 species (Klitgaard and Lavin 2005) 231taxa and cytotypes at four ploidy levels: Seeds of the putative hybrids were collected in the diploid (2x), tetraploid (4x), hexaploid (6x) and octoploid Municipio de la Huerta, Estado de Jalisco, Mexico, (8x) (Index to Plant Chromosome Numbers; Kawakami 19°29´N; 105°01´W (Carleial s/n, MEXU). The climate 1930; Bielig 1997; Arrighi et al. 2012, 2014; Chaintreuil is semi-dry and warm. Mean temperature in the area is et al. 2016, 2018; Brottier et al. 2018). America, where 25.2 °C, and there is a well-defined rainy season (average most of the taxa are 2n = 20 diploids, has been proposed annual precipitation: 1107 mm) occurring from June to as the center of origin of the genus, with a secondary October (García-Oliva et al. 2002). distribution in Africa and Asia where polyploid species The seeds of Aeschynomene evenia and A. scabra and some cases of aneuploidy predominate (Chaintreuil were collected in the municipalities of Coyuca de Cat- et al. 2018; Tapia-Pastrana et al. 2020). alán (18° 19´ N; 100° 42´ W, JC Soto 15333 (MEXU)) Although it is clear that in the Dalbergioid clade, and Arcelia (18°18´54´´N; 100°17´02´´W, JC Soto 15393 diploid 2n = 20 genera predominate, with some poly- (MEXU)) respectively, in the State of Guerrero, Mexi- ploid and aneuploid species, in Aeschynomene there is co. Both municipalities are part of the Tierra Caliente currently a renewed interest in knowing to what extent region. The predominant climate is warm subhumid polyploidy has contributed to the diversification and with rains from June to September (average annual radiation of the group. In this respect Arrighi et al. precipitation: 1100 to 1200 mm). The studied taxa are (2014) revealed multiple hybridization/polyploidiza- assigned to the infrageneric classification of Neotropi- tion events, highlighting the prominent role of allopoly- cal Aeschynomene sensu Rudd (1955) series Indicae of ploidy in the diversification of Nod-independent clade. subgenus Aeschynomene and are part of the Nod-inde- In addition Chaintreuil et al. (2016) studied African pendent monophyletic clade (Chaintreuil et al. 2013), Aeschynomene species and their data support the idea whose taxa are nodulated on roots and stems by photo- that the whole African group is fundamentally tetraploid synthetic Bradyrhizobium strains lacking the nod ABC and revealed the allopolyploid origin of A. afraspera genes necessary for the synthesis of Nod factors (Giraud et al. 2007). First cytogenetic register of an allopolyploid lineage of the genus Aeschynomene native to Mexico 19 2.2 Chromosome and karyotype procedures in putative of 20 individuals grown in pots under greenhouse con- hybrids ditions was evaluated. Interest was particularly focused on the number of leaflets and the presence of hairs on Seeds were collected in summer 2014 and from at their edges until the complete development of the fourth least six plants.
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