Centrophilic Retrotransposons of Plant Genomes

The centromeres of eukaryotic chromosomes are required to load CENH3/CENP-A variant nucleosomes and the kinetochore complex, which connects to spindle microtubules during cell division. Despite their conserved function, plant centromeres show rapid sequence evolution within and between species and a range of monocentric, holocentric, and polymetacentric architectures, which vary in kinetochore numbers and spacing. Plant centromeres are commonly composed of tandem satellite repeat arrays, which are invaded by specific families of centrophilic retrotransposons, whereas in some species the entire centromere is composed of such retrotransposons. We review the diversity of plant centrophilic retrotransposons and their mechanisms of integration, together with how epigenetic information and small RNAs control their proliferation. We discuss models for rapid centromere sequence evolution and speculate on the roles that centrophilic retrotransposons may play in centromere dynamics. We focus on plants but draw comparisons with animal and fungal centromeric transposons to highlight conserved and divergent themes across the eukaryotes.