Intracellular gene transfer and codon usage of cytoplasmic genomes in green plants

Intracellular gene transfer is widely recognized as one of the most important driving forces for species evolution. Here we investigated transferred cytoplasmic motifs in green plants including spore-bearing plants and seed-bearing plants (hereafter termed spore plants and seed plants) . Our analyses revealed that gene transfer in spore plants was characterized by shorter motifs than that of seed plants. Several spore species did not exhibit intracellular gene transfer. Meanwhile, high frequency transferred tRNA genes were identified with average values of minimum free energy at moderate level. From the chloroplast to the mitochondrial genome, trnP was found to have transferred with high frequency in green plants. In gene transfer from the mitochondrial to the chloroplast genome, trnN was found to be a highly transferred gene. We observed that several tRNA genes including trnF, trnW, and trnN were involved in bidirectional transfer, which may be related to application strategy of functional protein-coding genes in a plant's adaptive evolution. Codon Adaptation Index (CAI) analysis showed that codon usage was unbalanced in spore and seed plants. CAI values for seed plants were higher than those for spore plants in general, which may reveal rapid divergence adaptability of codon usage in the former. These results provide novel insights into gene transfer and codon usage within cytoplasmic genomes.