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1.
Reprod Domest Anim ; 2021 Mar 15.
Artigo em Inglês | MEDLINE | ID: mdl-33720451

RESUMO

The aim of this study was to analyse the association between single-nucleotide polymorphisms within INHA and ACVR2B and litter size in Dazu black goats. In total, twenty-two SNPs were genotyped in 190 individuals by SNaPshot and resequencing. The results showed that three SNPs (SNP_1, SNP_12 and SNP_13 in this study) were detected to have significant additive genetic effect on the recorded goat litter size (p < .05). The SNP_1 (NC_030809.1), a non-synonymous substitution of G for T at chr2-g. 28314990 in the exon 2 of INHA gene (NM_001285606.1), resulted in homozygote 2 (HOM2) contributed 0.25 and heterozygote (HET) contributed 0.12 larger litter than homozygote 1 (HOM1). Meanwhile, SNP_12 (Chr22-g. 11721225 A > T) and SNP_13 (Chr22-g. 11721227 A > C) (NC_030829.1) simultaneously mutated at the first and third position of a triplet AAA (lysine, K) in the exon 4 of ACVR2B gene (XM_018066623.1) had estimated genetic effects of HOM1 (0.00) and HOM2 (0.03) larger than HET (-0.12). In conclusion, one SNPs (chr2-g. 28314990 T > G) within the exon 2 of INHA and two SNPs (Chr22-g. 11721225 A > T and Chr22-g. 11721227 A > C) in the exon 4 of ACVR2B gene were highly recommended as candidate markers of litter size in Dazu black goats. A large-scale association study to assess the impact of these variants on litter size is still necessary.

2.
Anim Biotechnol ; : 1-19, 2021 Feb 26.
Artigo em Inglês | MEDLINE | ID: mdl-33635178

RESUMO

The genetic diversity and population structures of five Chongqing local chicken populations were investigated using by 24 microsatellite markers. Results revealed that the mean number of alleles (NA) ranged from 7.08 (Daninghe chicken, DN) to 8.46 (Nanchuan chicken, NC). The highest observed heterozygosity (HO) and expected heterozygosity (HE) were observed in DN (HO = 0.7252; HE = 0.7409) and the lowest HO and HE were observed in XS (Xiushan native chicken [XS], HO = 0.5910 and HE = 0.6697). The inbreeding coefficient (FIS) within population ranged from 0.022 (DN) to 0.119 (XS). Among the 24 microsatellite markers, four loci (MCW0111, MCW0016, ADL0278, and MCW0104) deviated from the Hardy-Weinberg equilibrium in all the studied populations. The results of population polygenetic analysis based on Nei's genetic distance and STRUCTURE software showed that the clustering of the five populations was incomplete consistent with geographical distribution. Moreover, a large number of gene flows were widespread among different populations, suggesting that genetic material exchanges occurred due to human activities and migration which was also verified by PCoA. In summary, this study preliminarily showed that Chongqing local chicken populations had rich genetic diversity and remarkable genetic divergence, but still high risk in conversion. These findings would be useful to the management of conservation strategies and the utilization of local chicken populations in further.

3.
Mol Immunol ; 124: 83-90, 2020 08.
Artigo em Inglês | MEDLINE | ID: mdl-32544655

RESUMO

Major histocompatibility complex (MHC) genes are critical for disease resistance or susceptibility responsible for host-pathogen interactions determined mainly by extensive polymorphisms in the MHC genes. Here, we examined the diversity and phylogenetic pattern of MHC haplotypes reconstructed using three MHC-linked microsatellite markers in 55 populations of five Bovidae species and compared them with those based on neutral autosomal microsatellite markers (NAMs). Three-hundred-and-forty MHC haplotypes were identified in 1453 Bovidae individuals, suggesting significantly higher polymorphism and heterozygosity compared with those based on NAMs. The ambitious boundaries in population differentiation (phylogenetic network, pairwise FST and STRUCTURE analyses) within and between species assessed using the MHC haplotypes were different from those revealed by NAMs associated closely with speciation, geographical distribution, domestication and management histories. In addition, the mean FST was significantly correlated negatively with the number of observed alleles (NA), observed (HO) and expected (HE) heterozygosity and polymorphism information content (PIC) (P < 0.05) in the MHC haplotype dataset while there was no correction of the mean FST estimates (P> 0.05) between the MHC haplotype and NAMs datasets. Analysis of molecular variance (AMOVA) revealed a lower percentage of total variance (PTV) between species/groups based on the MHC-linked microsatellites than NAMs. Therefore, it was inferred that individuals within populations accumulated as many MHC variants as possible to increase their heterozygosity and thus the survival rate of their affiliated populations and species, which eventually reduced population differentiation and thereby complicated their classification and phylogenetic relationship inference. In summary, host-pathogen coevolution and heterozygote advantage, rather than demographic history, act as key driving forces shaping the MHC diversity within the populations and determining the interspecific MHC diversity.


Assuntos
Animais Domésticos/genética , Evolução Biológica , Interações Hospedeiro-Patógeno/genética , Complexo Principal de Histocompatibilidade/genética , Animais , Animais Domésticos/imunologia , Bovinos , Variação Genética , Haplótipos , Heterozigoto , Interações Hospedeiro-Patógeno/imunologia , Complexo Principal de Histocompatibilidade/imunologia , Repetições de Microssatélites , Filogenia
4.
Anim Biotechnol ; : 1-6, 2020 Apr 15.
Artigo em Inglês | MEDLINE | ID: mdl-32293982

RESUMO

This study aims to identify the relative Copy number variation (CNV) associated with the litter size of Dazu black goats based on the unpublished CNV analytical results of our previously published sequencing data, in which the litter-size groups were classified into extreme low- and high-yield groups. Firstly, to compare the existence of valuable CNV in Dazu black goats with different fertility levels with mixed pools. We obtained 4992 and 4888 CNVs from the HY and LY, which overlapping 1461 genes, and classified on the original CNV type. Three genes [LOC108633278, PPP1R12A, and YIPF4] were observed in the intersection between the HY deletion and the LY duplication groups. Secondly, on individuals level, we identified a novel candidate CNV (Chr1_50215501, FST = 0.148, VST = 0.347) from 214 autosomal credible CNVs to be significant with litter size in the Dazu black goat, which located in the CBLB gene. This finding indicates the CBLB gene may affect the litter size of the Dazu black goats through structural variations, and Chr1_50215501 can be an effective genetic marker for marker-assisted selection breeding, and this study was also helps understand the molecular mechanism related to the goat litter size.

6.
3 Biotech ; 9(9): 336, 2019 Sep.
Artigo em Inglês | MEDLINE | ID: mdl-31475088

RESUMO

Litter size is considered to be the most important index for estimating domestic animal productivity. The number of indigenous goats in China with higher litter sizes than those of commercial breeds in other countries may be helpful for accelerating genetic improvements in goat breeding. We performed a genome-wide selective sweep analysis of 31 Dazu black goats with extreme standard deviation in litter size within the third fetus to identify significant genomic regions and candidate genes through different analyses. The analysis identified a total of 33,917,703 variants, including 32,262,179 SNPs and 1,655,524 indels. In addition, two novel candidate genes (LRP1B and GLRB), which are related to litter size, were obtained with π, Tajima's D, πA/πB, and F ST at the individual level with a 95% threshold for each parameter. These two genes were annotated in five GO terms (localization, binding, macromolecular complex, membrane part, and membrane) and two pathways (long-term depression and neuroactive ligand-receptor interaction pathway). Regarding the result of linkage disequilibrium (LD) analysis, in LRP1B and GRID2, the high-yield Dazu black goats exhibit significantly different LD patterns from low-yield goats. Litter size variability has low heritability and is related to multiple complex factors found in domestic animals. Obtaining a clear explanation and significant signal by genome-wide selective sweep analysis with a small sample size is difficult. However, we investigated some candidate genes, particularly LRP1B and GLRB, which may provide useful information for further research.

8.
Ecol Evol ; 9(10): 5916-5924, 2019 May.
Artigo em Inglês | MEDLINE | ID: mdl-31161008

RESUMO

Background: To assess the genetic diversity of seven Yunnan indigenous goat populations (Fengqing hornless goat, Mile red-bone goat, Longling goat, Ninglang black goat, Black-bone goat, Yunling black goat, and Zhaotong goat), their population structures were investigated using 20 microsatellite markers. Results: The results indicated that the genetic diversity of these goats was rich. The observed heterozygosity ranged from 0.4667 ± 0.0243 to 0.5793 ± 0.0230, and the mean number of alleles ranged from 4.80 ± 1.61 and 4.80 ± 1.64 to 6.20 ± 2.93. The population structure analysis showed that these seven goat populations were separated into two clusters, consistent with the results from phylogenetic networks, pairwise differences, and STRUCTURE analyses. We speculate that this may have been caused by natural geographical isolation, human migration and economic and cultural exchanges. We suggest removing CSRD247 and ILSTS005, two loci identified to be under positive selection in the present study, from the microsatellite evaluation system of goats. Conclusions: The present study may provide a scientific basis for the conservation and utilization of Yunnan indigenous goats.

9.
Mol Biol Rep ; 46(5): 5517-5523, 2019 Oct.
Artigo em Inglês | MEDLINE | ID: mdl-31175513

RESUMO

Litter size is considered to be the most important index for estimating domestic animal productivity. Due to its complexity, the molecular mechanism of litter size has not been elucidated, and it has restricted the use of marker-assisted selection to create high-yield populations in goats. A genome-wide selective sweep analysis was performed with 31 Dazu black goats to identify significant genomic regions and candidate genes related to litter size by a mixed pools strategy. A total of 96 candidate genes were identified, including NR6A1, STK3, IGF2BP2, AR, HMGA2, NPTX1, ANKRD17, DPYD, CLRB, PPP3CA, PLCB1, STK3 and HMGA2, using mixed pool analysis with ZHp and FST. We classified these candidate genes based on the functional classification and annotation of signaling pathways. According to the GO and KEGG analysis results, a total of 43 GO terms and 108 pathways were annotated from these genes. In particular, some novel candidate genes were enriched in reproduction-related pathways, such as the estrogen signaling pathway and oocyte meiosis. These findings provide insight into the influences of coding genes on the fecundity traits of goats.


Assuntos
Fertilidade/genética , Cabras/genética , Tamanho da Ninhada de Vivíparos/genética , Criação de Animais Domésticos/métodos , Animais , Genoma , Genômica , Genótipo , Fenótipo , Sequenciamento Completo do Genoma/métodos
10.
J Anim Breed Genet ; 136(5): 371-377, 2019 Sep.
Artigo em Inglês | MEDLINE | ID: mdl-31062447

RESUMO

The domestic yak (Bos grunniens) is an iconic symbol of animal husbandry on the Qinghai-Tibet Plateau. Long-term domestication and natural selection have led to a wide distribution of yak, forming many ecological populations to adapt to the local ecological environment. High altitude is closely related to oxygen density, and it is an important environmental ecological factor for biological survival and livestock production. The aim of the present study was to perform a preliminary analysis to identify the candidate genes of altitude distribution adapted ecological thresholds in yak using next-generation sequence technology. A total of 15,762,829 SNPs were obtained from 29 yaks with high- and low-altitude distribution by genome-wide sequencing. According to the results of the selective sweep analysis with FST and ZHp, 21 candidate genes were identified. 14 genes (serine/threonine protein kinase TNNI3K, TEN1, DYM, ITPR1, ZC4H2, KNTC1, ADGRB3, CLYBL, TANGO6, ASCC3, KLHL3, PDE4D, DEPDC1B and AGBL4) were grouped into 32 Gene Ontology terms, and four genes (RPS6KA6, ITPR1, GNAO1 and PDE4D) annotated in 35 pathways, including seven environmental information processing and one environmental adaptation. Therefore, the novel candidate genes found in the current study do not only support new theories about high-altitude adaptation, but also further explain the molecular mechanisms of altitude adaptation threshold in yaks.


Assuntos
Bovinos/genética , Bovinos/fisiologia , Estudo de Associação Genômica Ampla , Adaptação Biológica , Altitude , Animais , Polimorfismo de Nucleotídeo Único
11.
3 Biotech ; 9(3): 99, 2019 Mar.
Artigo em Inglês | MEDLINE | ID: mdl-30800610

RESUMO

Sex reversal has been studied extensively in vertebrate species, particularly in domestic goats, because polled intersex syndrome (PIS) has seriously affected their production efficiency. In the present study, we used histopathologically diagnosed cases of PIS to identify correlated genomic regions and variants using representative selection signatures and performed GWAS using Restriction-Site Associated Resequencing DNA. We identified 171 single-nucleotide polymorphisms (SNPs) that may have contributed to this phenotype, and 53 SNPs were determined to be located in coding regions using a general linear model. The transcriptome data sets of differentially expressed genes (DEGs) in the pituitary tissues of intersexual and nonintersexual goats were examined using high-throughput technology. A total of 10,063 DEGs and 337 long noncoding RNAs were identified. The DEGs were clustered into 56 GO categories and determined to be significantly enriched in 53 signaling pathways by KEGG analysis. In addition, according to qPCR results, PSPO2 and FSH were significantly more highly expressed in sexually mature pituitary tissues of intersexual goats compared to healthy controls (nonintersexual). These results demonstrate that certain novel potential genomic regions may be responsible for intersexual goats, and the transcriptome data indicate that the regulation of various physiological systems is involved in intersexual goat development. Therefore, these results provide helpful data for understanding the molecular mechanisms of intersex syndrome in goats.

12.
Mol Biol Rep ; 46(1): 1307-1311, 2019 Feb.
Artigo em Inglês | MEDLINE | ID: mdl-30560407

RESUMO

China has numerous local domestic sheep breeds. In this study, the genetic diversity of eight sheep populations was estimated using 17 microsatellites. Knowledge of such diversity provides novel insight into the degree of breed protection needed and the prediction of hybrid advantage. In total, 17 microsatellites were genotyped in 186 individuals from eight populations. The mean number of alleles (± SD) ranged from 3.71 ± 1.36 in Zhaotong sheep to 11.94 ± 3.58 in small-tailed Han sheep. The observed heterozygote frequency (± SD) within a population ranged from 0.482 ± 0.025 in Zhaotong sheep to 0.664 ± 0.023 in Tibetan sheep. In addition, using pairwise difference (FST) analysis, the highest within-population diversity was observed in Tibetan sheep (πX = 12.8098) and small-tailed Han (πX = 12.67873), and the lowest diversity was observed in Zhaotong sheep (πX = 7.90337). The results for genetic divergence between populations indicated that the populations were significantly different (P < 0.05) based on the average number of pairwise differences between populations (πXY) and the corrected average pairwise differences. Both phylogenetic networks and structure analysis showed that these eight populations were separated into three clusters in accordance with their geographical habitat, except Tibetan and Hu sheep. In short, we genotyped eight local Chinese sheep populations using 17 microsatellites, and the results indicated that their current genetic diversity is decreasing and that new conservation strategies are needed. In addition, significant genetic differences between populations could be used in cross breeding.


Assuntos
Variação Genética , Genética Populacional , Ovinos/genética , Animais , Cruzamento , China , Filogenia
13.
Ecol Evol ; 8(10): 5111-5123, 2018 May.
Artigo em Inglês | MEDLINE | ID: mdl-29876086

RESUMO

The objective of this study was to assess the genetic diversity and population structure of goats in the Yangtze River region using microsatellite and mtDNA to better understand the current status of those goat genetic diversity and the effects of natural landscape in fashion of domestic animal genetic diversity. The genetic variability of 16 goat populations in the littoral zone of the Yangtze River was estimated using 21 autosomal microsatellites, which revealed high diversity and genetic population clustering with a dispersed geographical distribution. A phylogenetic analysis of the mitochondrial D-loop region (482 bp) was conducted in 494 goats from the Yangtze River region. In total, 117 SNPs were reconstructed, and 173 haplotypes were identified, 94.5% of which belonged to lineages A and B. Lineages C, D, and G had lower frequencies (5.2%), and lineage F haplotypes were undetected. Several high-frequency haplotypes were shared by different ecogeographically distributed populations, and the close phylogenetic relationships among certain low-frequency haplotypes indicated the historical exchange of genetic material among these populations. In particular, the lineage G haplotype suggests that some west Asian goat genetic material may have been transferred to China via Muslim migration.

14.
Artigo em Inglês | MEDLINE | ID: mdl-27841052

RESUMO

The mitochondrial DNA control region (D-loop) is a widely used molecular marker in evolutionary and phylogeographic research. However, the occurrence of heteroplasmy of the D-loop region within individuals has rarely been investigated. In this study, a total of 85 Chinese sheep were used to amplify a partial D-loop region, and 15 heteroplasmic animals (17.64%) were identified. A comparative analysis of the PCR amplification and cloning of the D-loop sequences from the heteroplasmic samples revealed most of the sequencing profile from the heteroplasmic regions started at the beginning of a 75-bp random repeat motif. In addition, a total of 22 nonsyngeneic sequences with a D-loop were found in 61 of the clones obtained from the 4 random heteroplasmic and 3 homozygote animals, and their genomic locations were compared for homology. In summary, the D-Loop sequencing profiles appear to be heteroplasmic and could arise from tandem repeat motifs and unspecific replication during PCR amplification; however, they are not likely due to the presence of multiple mitochondrial genomes within an individual.


Assuntos
Variação Genética , Genoma Mitocondrial , Mitocôndrias/genética , Ovinos/genética , Sequências de Repetição em Tandem , Animais , Filogenia , Sequências Reguladoras de Ácido Nucleico
15.
Ecol Evol ; 6(3): 810-7, 2016 Feb.
Artigo em Inglês | MEDLINE | ID: mdl-26865968

RESUMO

The domestic sheep (Ovis aries) has been an economically and culturally important farm animal species since its domestication around the world. A wide array of sheep breeds with abundant phenotypic diversity exists including domestication and selection as well as the indigenous breeds may harbor specific features as a result of adaptation to their environment. The objective of this study was to investigate the population structure of indigenous sheep in a large geographic location of the Chinese mainland. Six microsatellites were genotyped for 611 individuals from 14 populations. The mean number of alleles (±SD) ranged from 7.00 ± 3.69 in Gangba sheep to 10.50 ± 4.23 in Tibetan sheep. The observed heterozygote frequency (±SD) within a population ranged from 0.58 ± 0.03 in Gangba sheep to 0.71 ± 0.03 in Zazakh sheep and Minxian black fur sheep. In addition, there was a low pairwise difference among the Minxian black fur sheep, Mongolian sheep, Gansu alpine merino, and Lanzhou fat-tailed sheep. Bayesian analysis with the program STRUCTURE showed support for 3 clusters, revealing a vague genetic clustering pattern with geographic location. The results of the current study inferred high genetic diversity within these native sheep in the Chinese mainland.

16.
Mitochondrial DNA A DNA Mapp Seq Anal ; 27(6): 3826-3827, 2016 11.
Artigo em Inglês | MEDLINE | ID: mdl-25186658

RESUMO

The Chinese Yakow, Bos primigenius taurus × Bos grunniens, is a large and commercially important hybrid in family Bovidae. We first determined and annotated its complete mitochondrial genome. The mitogenome is 16,322 bp in length, consisting of 13 protein-coding genes, 22 transfer RNA (tRNA) genes, 2 ribosomal RNA (rRNA) genes and a control region. As in other mammals, most mitochondrial genes are encoded on the heavy strand, except for ND6 and eight tRNA genes, which are encoded on the light strand. Its overall base composition is A: 33.7%, T: 27.3%, C: 25.8% and G: 13.2%. The complete mitogenome of Yakow (B. p. taurus × B. grunniens) could provide an important data to further explore the taxonomic status of Yakow (B. p. taurus × B. grunniens) in B. grunniens and enrich the genetic information or evolutionary history of the Bovidae.


Assuntos
Bovinos/genética , Genoma Mitocondrial/genética , Animais , Composição de Bases , Sequência de Bases , Bovinos/classificação , China , DNA Mitocondrial/química , DNA Mitocondrial/genética , Proteínas Mitocondriais/genética , Fases de Leitura Aberta/genética , RNA Ribossômico/genética , RNA de Transferência/genética , Análise de Sequência de DNA
17.
Mitochondrial DNA A DNA Mapp Seq Anal ; 27(5): 3171-2, 2016 09.
Artigo em Inglês | MEDLINE | ID: mdl-25731719

RESUMO

Dazu Black goat is an indigenous goat genetic resource in Southwest of China. Here, we describe its complete mitochondrial genome sequence. The mitogenome is 16,641 bp in length, consisting of 13 protein-coding genes, 22 transfer RNA (tRNA) genes, 2 ribosomal RNA (rRNA) genes and a control region. As in other mammals, most mitochondrial genes are encoded on the heavy strand, except for ND6 and eight tRNA genes, which are encoded on the light strand. Its overall base composition is A: 33.5%, T: 27.3%, C: 26.1% and G: 13.1%. The complete mitogenome of the indigenous goat could provide important data to further explore the taxonomic status of the subspecies and also provide a starting point for further phylogenetic studies.


Assuntos
Genoma Mitocondrial , Cabras/genética , Animais , Composição de Bases , NADH Desidrogenase/genética , Fases de Leitura Aberta , Filogenia , RNA Ribossômico/genética , RNA de Transferência/genética
18.
Mitochondrial DNA A DNA Mapp Seq Anal ; 27(5): 3308-9, 2016 09.
Artigo em Inglês | MEDLINE | ID: mdl-25693713

RESUMO

Here, we describe the complete mitochondrial genome sequences of Jining Gray goat, Fushun black goat, Youzhou black-skin goat, and Hechuan white goat. The mitogenome of those four goats consisted of 16,640 nt, consisting of 13 protein-coding genes, 22 transfer RNA (tRNA) genes, 2 ribosomal RNA (rRNA) genes and a control region. As in other mammals, most mitochondrial genes are encoded on the heavy strand, except for ND6 and eight tRNA genes, which are encoded on the light strand. The complete mitogenome of these four local breeds of Chinese native goats could provide an important data to further breed improvement and animal genetics resource conservation in China.


Assuntos
Genoma Mitocondrial , Cabras/genética , Animais , DNA Intergênico/química , DNA Intergênico/genética , Fases de Leitura Aberta , RNA Ribossômico/genética , RNA de Transferência/genética
19.
Mitochondrial DNA A DNA Mapp Seq Anal ; 27(5): 3063-4, 2016 09.
Artigo em Inglês | MEDLINE | ID: mdl-25740215

RESUMO

The Chuanzhong black goat (Capra hircus) is a breed native to southwest of China. Its complete mitochondrial genome is 16,641 nt in length, consisting of 13 protein-coding genes, 22 transfer RNA (tRNA) genes, two ribosomal RNA (rRNA) genes, and a non-coding control region. As in other mammals, most mitochondrial genes are encoded on the heavy strand, except for ND6 and eight tRNA genes, which are encoded on the light strand. Its overall base composition is A: 33.5%, T: 27.3%, C: 26.1%, and G: 13.1%. The complete mitogenome of the Chinese indigenous breed of goat could provide a basic data for further phylogenetics analysis.


Assuntos
Genoma Mitocondrial , Cabras/classificação , Cabras/genética , Animais , Composição de Bases , Cruzamento , China , Genes Mitocondriais , Tamanho do Genoma , Fases de Leitura Aberta , Filogenia , Análise de Sequência de DNA , Sequenciamento Completo do Genoma
20.
Vet Immunol Immunopathol ; 167(3-4): 196-9, 2015 Oct 15.
Artigo em Inglês | MEDLINE | ID: mdl-26257111

RESUMO

In this study, the genetic variability of Chinese indigenous goat breeds (Capra hircus) was analyzed using the MHC-associated microsatellite markers BF1, BM1818, BM1258, and DYMS1. To examine genetic variability, the levels of heterozigosity, degrees of inbreeding, and genetic differences among the breeds were analyzed. The mean number of alleles ranged from 5.50±3.70 in Enshi black goats (EB) to 11.50±3.70 in the Jianyang big ear (JE) breed. The mean observed heterozygosity and mean expected heterozygosity varied from 0.25±0.04 in Jining Qing goats (JQ) to 0.54±0.05 in Chuannan black goats (CN) and from 0.49±0.18 in Hechuan white goats (HW) to 0.78±0.05 in JE, respectively. The mean FIS values ranged from 0.23 in HW to 0.51 in JQ. In addition, the genetic variation among populations and geographic location did indicate a correlation of genetic differences with geographic distance, which was revealed by the phylogenetic network. In conclusion, the high variability and population structure among Chinese native goats in the Major Histocompatibility Complex would be caused by co-evolution between MHC alleles and the epidemic history or pathogens in different agro-ecological zones.


Assuntos
Cabras/genética , Cabras/imunologia , Complexo Principal de Histocompatibilidade , Alelos , Animais , Cruzamento , China , Evolução Molecular , Variação Genética , Genética Populacional , Heterozigoto , Repetições de Microssatélites , Filogenia , Polimorfismo Genético
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