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Intrapopulation Chromosomal Polymorphism in Mazama Gouazoubira (Cetartiodactyla; Cervidae): the Emergence of a New Species?

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Intrapopulation Chromosomal Polymorphism in Mazama Gouazoubira (Cetartiodactyla; Cervidae): the Emergence of a New Species? Original Article Cytogenet Genome Res 2018;154:147–152 Accepted: February 22, 2018 DOI: 10.1159/000488377 by M. Schmid Published online: April 14, 2018 Intrapopulation Chromosomal Polymorphism in Mazama gouazoubira (Cetartiodactyla; Cervidae): The Emergence of a New Species? Mirela P. Valeri Iara M. Tomazella José M.B. Duarte Núcleo de Pesquisa e Conservação de Cervídeos (NUPECCE), Departamento de Zootecnia, Faculdade de Ciências Agrárias e Veterinárias, Universidade Estadual Paulista (UNESP), Jaboticabal , Brazil Keywords vantage. On the other hand, these polymorphisms may neg- B chromosome · Centric fusion · G-banding · Gray brocket atively influence fertility and raise questions about sustain- deer ability or reproductive isolation of the population. © 2018 S. Karger AG, Basel Abstract Mazama gouazoubira is a small deer species widely distrib- Mazama gouazoubira , the gray or brown brocket deer, uted in South America. Previous studies have shown that this is a small deer species found from southern Uruguay to species presents intraspecific chromosomal polymorphisms, the north of the Mato Grosso state in Brazil and from the which could affect fertility due to the effects of chromosom- Andes to the Atlantic. Its habitat varies from closed Cer- al rearrangements on gamete formation. Important aspects rado to agricultural areas. This species can readily adapt regarding the karyotype evolution of this species and the to modified areas due to its high ecological plasticity [Du- genus remain undefined due to the lack of information con- arte, 2007; Rodrigues et al., 2014]. cerning the causes of this chromosomal variation. Nineteen M. gouazoubira (2n = 70; FN = 70) has 68 acrocentric individuals belonging to the Mazama gouazoubira popula- autosomes, an acrocentric X, and a metacentric Y [Neit- tion located in the Pantanal were cytogenetically evaluated. zel, 1987; Fontana and Rubini, 1990], similar to the puta- Among the individuals analyzed, 9 had B chromosomes and tive Cervidae ancestral karyotype [Dementyeva et al., 5 carried a heterozygous centric fusion (2n = 69 and FN = 70). 2010]. The karyotypes of other species in the Mazama In 3 individuals, the fusion occurred between chromosomes genus differ from the proposed ancestral deer karyotype X and 16, in 1 individual between chromosomes 7 and 21, by a high number of chromosomal rearrangements, in- and in another individual between chromosomes 4 and 16. cluding tandem fusions, centric fusions, pericentric in- These striking polymorphisms could be explained by several versions, and the presence of B chromosomes [Fontana hypotheses. One is that the chromosome rearrangements in and Rubini, 1990]. this species are recent and not fixed in the population yet, Considerable intraspecific variation in the chromo- and another hypothesis is that they represent a balanced some number exists in M. gouazoubira due to Robertso- polymorphism and that heterozygotes have an adaptive ad- nian translocations and B chromosomes [Duarte, 1992, © 2018 S. Karger AG, Basel José M.B. Duarte Departamento de Zootecnia, Faculdade de Ciências Agrárias e Veterinárias Universidade Estadual Paulista, Via de Acesso Prof. Paulo Donato Castellane, s/n E-Mail [email protected] Zona Rural, Jaboticabal, SP 14884–900 (Brazil) www.karger.com/cgr E-Mail barbanti @ fcav.unesp.br Downloaded by: UNESP Universidade Estdual Paulista 186.217.236.55 - 8/5/2019 7:07:37 PM 1998; Duarte and Jorge, 1996]. Some researchers hypoth- FN = 70, with 68 acrocentric autosomes, an acrocentric esize that this chromosome variability is due to chro- X, and a metacentric Y. The other 3 karyomorphs result- mosomal fragility, which promotes chromosomal breaks ed from Robertsonian rearrangements and presented and rearrangements [Duarte, 1998; Vargas-Munar et al., 2n = 69 and FN = 70 ( Fig. 1 ). If the Robertsonian rear- 2010]. Robertsonian rearrangements (centromeric fis- rangements in this population are fusions, the reduction sions and fusions) are the most commonly incorporated in the diploid number without altering the fundamental rearrangements in mammalian karyotype evolution. It is number is the result of a centric fusion between various known that this type of rearrangement can lower fitness acrocentric chromosomes forming a metacentric or sub- due to the production of unbalanced gametes [Baker and metacentric chromosome. These 3 karyomorphs differ Bickham, 1986]. Therefore, it would be important to bet- from each other because of the different chromosomes ter understand Robertsonian rearrangements due to their involved in the fusion. frequent occurrence in evolution and their influence on Of the 19 deer, 5 were carriers of a heterozygous Rob- fertility [Daniel, 1988; Ferlin et al., 2006]. Further, the ertsonian translocation; the chromosomes involved in mechanism about how Robertsonian rearrangements can the rearrangements were identified based on their G- become fixed in a population is not well understood. M. banding pattern. Among these 5 animals, 3 shared the gouazoubira populations are an ideal model for clarifying same karyotype in which the bi-armed chromosome, some of these issues. We know that the free-ranging classified as submetacentric, was the result of Robertso- Mazama population located in the Pantanal biome differs nian translocation between the X chromosome and auto- in the chromosome number due to Robertsonian rear- some 16 ( Fig. 1 A). The other 2 carriers of the Robertso- rangements. In this research, we identified the chromo- nian translocation showed bi-armed chromosomes, clas- somes involved in these rearrangements and tested the sified as metacentric. In 1 individual the translocation hypothesis that fusions are the responsible mechanism. We occurred between chromosomes 4 and 16 ( Fig. 1 B) and also provide data regarding their potential influence on fer- in the other between chromosomes 7 and 21 (Fig. 1 C). tility, fitness, and reproductive isolation of this population. Another source of chromosomal polymorphism iden- tified in the population was the inter- and intraindividu- al variation in dot-like B chromosomes, with 0–2 B chro- Materials and Methods mosomes present in addition to the basic complement in 9 individuals of the 19 deer analyzed ( Table 1 ). There was A major advantage of this research is that all animals studied no statistical difference (ANOVA test) in the presence of were wild-caught and the provenience is precisely known. Nine- teen (12 males and 7 females) free-ranging M. gouazoubira were B chromosomes between the individuals with a standard captured by drive net or darts with transmitters [Duarte et al., karyotype and the carriers of the Robertsonian transloca- 2010] within a 15 km radius in the Nhecolândia Pantanal in the tions. municipality of Corumbá, Mato Grosso do Sul, Brazil (19° 00′ 33′ S, 57° 39′ 12′ W). Skin biopsies were collected, frozen in liquid ni- trogen [Duarte et al., 1999], and later used to obtain chromosom- al preparations using fibroblast cultures [Verma and Babu, 1995] Discussion which were maintained until passage 4 [Magalhães et al., 2017]. Twenty metaphases of each specimen were analyzed to determine To our knowledge, this is the first cytogenetic study on the diploid and fundamental number. G-banding was performed a population of M. gouazoubira with all analyzed animals according to Seabright [1971] to correctly identify the chromo- caught in the wild in a restricted area. The observed in- somes involved in the rearrangements. The chromosomes were classified as metacentric, submetacentric, or acrocentric by arm traspecific chromosomal polymorphisms, such as the ratio [Levan et al., 1964]. B chromosomes were calculated sepa- presence of B chromosomes and Robertsonian transloca- rately of the diploid and fundamental number due to intraindi- tions, have also been reported for other species of the ge- vidual variation. nus Mazama: M. americana [Abril et al., 2010], M. nana [Abril and Duarte, 2008], M. bororo [Duarte and Jorge, 2003], M. nemorivaga [Duarte, 1998], and even for M. Results gouazoubira [Duarte, 1992], though not for such a re- stricted geographic region. In this work, fibroblast cul- Among the 19 deer analyzed, 4 different karyomorphs tures were maintained until passage 4, and according to were identified. The most frequent is considered the stan- Magalhães et al. [2017], M. gouazoubira fibroblast cell dard karyotype for the species and presented 2n = 70, lines remain viable and functional until passage 7. Thus, 148 Cytogenet Genome Res 2018;154:147–152 Valeri/Tomazella/Duarte DOI: 10.1159/000488377 Downloaded by: UNESP Universidade Estdual Paulista 186.217.236.55 - 8/5/2019 7:07:37 PM A B CD Fig. 1. G-banded karyotypes of M. gouazoubira . A Representative karyotype of the translocation between the X chromosome and autosome 16. B Representative karyotype of the translocation between autosomes 4 and 16. C Representative karyotype of the translocation between autosomes 7 and 21. D G-banded chromosomes and diagrammatic representation of the respective translocations. the observed chromosome polymorphism was not related fundamental number is centric fusion. Also, chromo- to the number of passages used. somal evolution in the Cervidae family is based on centric Further, for the first time in M. gouazoubira , we deter- and tandem fusions, leading to the reduction of the dip- mined which Robertsonian translocations were present loid number [Fontana and Rubini, 1990], as observed in within the population using G-banding. Since the chro- the species Muntiacus
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