Tc1/mariner and Zator, as two superfamilies of IS630-Tc1-mariner (ITm) group, have been well-defined. However, the molecular evolution and domestication of pogo transposons, once designated as an important family of the Tc1/mariner superfamily, are still poorly understood. Here, phylogenetic analysis show that pogo transposases, together with Tc1/mariner, DD34E/Gambol, and Zator transposases form four distinct monophyletic clades with high bootstrap supports (> = 74%), suggesting that they are separate superfamilies of ITm group. The pogo superfamily represents high diversity with six distinct families (Passer, Tigger, pogoR, Lemi, Mover, and Fot/Fot-like) and wide distribution with an expansion spanning across all the kingdoms of eukaryotes. It shows widespread occurrences in animals and fungi, but restricted taxonomic distribution in land plants. It has invaded almost all lineages of animals—even mammals—and has been domesticated repeatedly in vertebrates, with 12 genes, including centromere-associated protein B (CENPB), CENPB DNA-binding domain containing 1 (CENPBD1), Jrk helix–turn–helix protein (JRK), JRK like (JRKL), pogo transposable element derived with KRAB domain (POGK), and with ZNF domain (POGZ), and Tigger transposable element-derived 2 to 7 (TIGD2–7), deduced as originating from this superfamily. Two of them (JRKL and TIGD2) seem to have been co-domesticated, and the others represent independent domestication events. Four genes (TIGD3, TIGD4, TIGD5, and POGZ) tend to represent ancient domestications in vertebrates, while the others only emerge in mammals and seem to be domesticated recently. Significant structural variations including target site duplication (TSD) types and the DDE triad signatures (DD29–56D) were observed for pogo transposons. Most domesticated genes are derived from the complete transposase genes; but CENPB, POGK, and POGZ are chimeric genes fused with additional functional domains. This is the first report to systematically reveal the evolutionary profiles of the pogo transposons, suggesting that pogo and Tc1/Mariner are two separate superfamilies of ITm group, and demonstrating the repeated domestications of pogo in vertebrates. These data indicate that pogo transposons have played important roles in shaping the genome and gene evolution of fungi and animals. This study expands our understanding of the diversity of pogo transposons and updates the classification of ITm group.

中文翻译：

---

pogo（IS630-Tc1-mariner 转座子的一个独立超家族）的进化揭示了脊椎动物中反复出现的驯化事件。

Tc1/mariner 和 Zator 作为 IS630-Tc1-mariner (ITm) 组的两个超家族，已得到明确定义。然而，曾经被指定为Tc1/mariner超家族的重要家族的pogo转座子的分子进化和驯化仍然知之甚少。在这里，系统发育分析表明，pogo 转座酶与 Tc1/mariner、DD34E/Gambol 和 Zator 转座酶一起形成四个不同的单系进化枝，具有高引导支持 (> = 74%)，表明它们是 ITm 组的独立超家族。pogo 超家族代表着高度多样性，有 6 个不同的家族（Passer、Tigger、pogoR、Lemi、Mover 和 Fot/Fot-like），并且分布广泛，扩展到真核生物的所有王国。它在动物和真菌中广泛存在，但在陆地植物中的分类分布受到限制。它几乎入侵了所有动物谱系——甚至哺乳动物——并在脊椎动物中反复驯化，拥有12个基因，包括着丝粒相关蛋白B（CENPB）、CENPB DNA结合域含有1（CENPBD1）、Jrk螺旋-转-螺旋蛋白 (JRK)、JRK 样 (JRKL)、KRAB 结构域衍生的 pogo 转座元件 (POGK) 和 ZNF 结构域 (POGZ) 以及 Tigger 转座元件衍生的 2 至 7 (TIGD2–7)，推测源自这个超家族。其中两个（JRKL 和 TIGD2）似乎是共同驯化的，其他则代表独立的驯化事件。四个基因（TIGD3、TIGD4、TIGD5 和 POGZ）往往代表脊椎动物的古代驯化，而其他基因仅出现在哺乳动物中，并且似乎是最近被驯化的。pogo 转座子观察到显着的结构变化，包括靶位点重复 (TSD) 类型和 DDE 三联体特征 (DD29-56D)。大多数驯化基因源自完整的转座酶基因；但 CENPB、POGK 和 POGZ 是与其他功能域融合的嵌合基因。这是第一份系统揭示pogo转座子进化谱的报告，表明pogo和Tc1/Mariner是ITm组的两个独立的超家族，并证明了pogo在脊椎动物中的反复驯化。这些数据表明pogo转座子在塑造真菌和动物的基因组和基因进化方面发挥了重要作用。这项研究扩展了我们对 pogo 转座子多样性的理解，并更新了 ITm 组的分类。