*Review* **Nuclear Integrants of Organellar DNA Contribute to Genome Structure and Evolution in Plants**

**Guo-Jun Zhang 1,2, Ran Dong 1, Li-Na Lan 1, Shu-Fen Li 1, Wu-Jun Gao 1,\* and Hong-Xing Niu 1,\***


Received: 27 December 2019; Accepted: 18 January 2020; Published: 21 January 2020

**Abstract:** The transfer of genetic material from the mitochondria and plastid to the nucleus gives rise to nuclear integrants of mitochondrial DNA (NUMTs) and nuclear integrants of plastid DNA (NUPTs). This frequently occurring DNA transfer is ongoing and has important evolutionary implications. In this review, based on previous studies and the analysis of NUMT/NUPT insertions of more than 200 sequenced plant genomes, we analyzed and summarized the general features of NUMTs/NUPTs and highlighted the genetic consequence of organellar DNA insertions. The statistics of organellar DNA integrants among various plant genomes revealed that organellar DNA-derived sequence content is positively correlated with the nuclear genome size. After integration, the nuclear organellar DNA could undergo different fates, including elimination, mutation, rearrangement, fragmentation, and proliferation. The integrated organellar DNAs play important roles in increasing genetic diversity, promoting gene and genome evolution, and are involved in sex chromosome evolution in dioecious plants. The integrating mechanisms, involving non-homologous end joining at double-strand breaks were also discussed.

**Keywords:** genome structure; genome evolution; DNA transfer; nuclear integrants of plastid DNA (NUPT); nuclear integrants of mitochondrial DNA (NUMT)
