plant mitochondria

Kubo T & Newton KJ 2008 Angiosperm mitochondrial genomes and mutations. Mitochondrion 8:5-14.

  • the sizes of angiosperm mitochondrial genomes have expanded since plants colonized the land although the genomes have lost some genes during this evolutionary period
  • intergenic regions have expanded in angiosperm mitochondria
  • the evolutionary trend of the angiosperm mitochondrial genome is counter to that of the mammalian mitochondrial genome
  • another characteristic of angiosperm mitochondrial genomes is the high degree of variation in terms of the genomic organization
  • frequent rearrangements are still ongoing in almost every angiosperm lineage
  • the complete nucleotide sequences of 15 mitochondrial genomes from seven angiosperm species have been made available (Table 1)
  • the sizes of the master chromosomes of the sequenced mitochondrial genomes range from 221,853 (rapeseed) to 739,719 bp (maize CMS-C; Fig. 1)
  • the number of mitochondrial genes in angiosperms is 50–60 (not considering copy number)
  • some of the mitochondrial genes in angiosperms are interrupted by introns
  • in each of the sequenced genomes, the total number of the introns is 20–24, constituting 4–13% of the genome
  • all the introns in the sequenced mitochondrial genomes are classified as group II type
  • a horizontally transferred group I intron has also been documented
  • one of the characteristics of angiosperm mitochondrial introns is the presence of trans splicing, particularly for genes encoding Complex I subunits
  • RNA editing is a necessary step for angiosperm mitochondrial gene expression, which converts cytidine residues to uridine and, rarely, uridine to cytidine
  • most of the RNA editing sites are found in the protein-coding genes, and a few are in tRNAs, untranslated regions, and introns (Shikanai, 2006)
  • the numbers of editing sites have been determined for some of the sequenced angiosperm species
  • a total of 357 in sugar beet
  • 491 in rice
  • the gene-coding regions constitute only 7–17% of the mitochondrial genome in the angiosperms sequenced to date
  • the remaining regions (intergenic regions) were examined to discover how the genome sizes had expanded in angiosperms
  • no conclusive mechanism has been proposed yet
  • some intergenic regions contain chloroplast DNA sequences (1.6–6.2% of the genome) and nuclear DNA sequences (0.1–13.4% of the genome)
  • the arrangement of mitochondrial genes can vary, even within a single angiosperm species
  • angiosperm mitochondrial genomes have undergone frequent genome rearrangement
  • rearrangement points are unevenly distributed in the genome
  • the reason for this is unclear
  • most of the rearrangements do not affect gene expression
  • they just produce a polymorphism between the genomes
  • in rare cases they are critical for mitochondrial gene expression
  • in angiosperms, the most commonly seen class of mitochondrial mutant is cytoplasmic male sterility
  • efforts to identify the responsible genes in mitochondrial genomes resulted in the discovery of aberrant open reading frames, consisting of fragments of mitochondrial genes and/or unknown sequences, usually with a chimeric structure
  • in a few cases, aberrant ORFs are translated into unique polypeptides associated with mutant phenotypes
  • aberrant ORFs can also exist within gene-coding regions such as the 5' leader of the atp6 gene
  • a number of aberrant ORFs, each producing unique polypeptides, have been reported to be associated with CMS
  • the aberrant ORFs have little in common, even when their phenotype is very similar
  • the CMS-associated aberrant ORFs have arisen independently during the course of angiosperm evolution
  • accumulation of CMS-associated proteins may affect all the tissues but that there is a mechanism of compensation in non-anther tissues
  • CMS-associated aberrant ORFs often neighbor, and are co-transcribed together with genes for ATPase subunits
  • from an evolutionary viewpoint, comparative analyses of sequenced mitochondrial genomes revealed that flanking regions of the genes for ATPase subunits are less well conserved compared to those of other genes
  • suggesting that mutations can accumulate more readily in these regions