Germline-restricted chromosomes of the songbirds

P. M. Borodin
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Abstract

Germline-restricted chromosomes (GRCs) are present in the genomes of germline cells and absent from somatic cells. A GRC is found in all species of the songbirds (Passeri) and in none of the other bird orders studied to date. This indicates that GRC originated in the common ancestor of the songbirds. The germline-restricted chromosome is permanently absent from somatic cells of the songbird, while female germline cells usually contain two copies of GRC and male ones have one copy. In females, GRCs undergo synapsis and restricted recombination in their terminal regions during meiotic prophase. In males, it is almost always eliminated from spermatocytes. Thus, GRC is inherited almost exclusively through the maternal lineage. The germline-restricted chromosome is a necessary genomic element in the germline cells of songbirds. To date, the GRC genetic composition has been studied in four species only. Some GRC genes are actively expressed in female and male gonads, controlling the development of germline cells and synthesis of the proteins involved in the organization of meiotic chromosomes. Songbird species vary in GRC size and genetic composition. The GRC of each bird species consists of amplified and modified copies of genes from the basic genome of that species. The level of homology between GRCs of different species is relatively low, indicating a high rate of genetic evolution of this chromosome. Transmission through the maternal lineage and suppression of the recombination contribute significantly to the accelerated evolution of GRCs. One may suggest that the rapid coordinated evolution between the GRC genes and the genes of the basic genome in the songbirds might be responsible for the explosive speciation and adaptive radiation of this most species-rich and diverse infraorder of birds.
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鸣禽的生殖系限制染色体
生殖系限制性染色体(GRCs)存在于生殖系细胞的基因组中,而不存在于体细胞中。在所有鸣禽(雀鸟)中都发现了GRC,而在迄今为止所研究的其他鸟类中没有发现GRC。这表明GRC起源于鸣禽的共同祖先。鸣禽的体细胞中永远不存在生殖系限制性染色体,而雌性生殖系细胞通常含有两个GRC拷贝,雄性生殖系细胞只有一个GRC拷贝。在雌性中,GRCs在减数分裂前期在其末端区域发生突触和限制性重组。在男性中,它几乎总是从精母细胞中消除。因此,GRC几乎完全通过母系遗传。生殖系限制性染色体是鸣禽生殖系细胞中必不可少的基因组元件。迄今为止,仅在四个物种中研究了GRC的遗传组成。一些GRC基因在雌性和雄性性腺中活跃表达,控制生殖细胞的发育和参与减数分裂染色体组织的蛋白质的合成。鸣禽的GRC大小和基因组成各不相同。每种鸟类的GRC由该物种基本基因组的扩增和修饰基因拷贝组成。不同物种GRCs间同源性较低,表明该染色体具有较高的遗传进化速率。通过母系传播和抑制重组显著促进了GRCs的加速进化。鸣禽的GRC基因与基本基因组基因之间的快速协调进化可能是这种鸟类中物种最丰富和最多样化的次目物种形成和适应性辐射爆发的原因。
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