We have cloned and sequenced the genes atpB and atpE, coding for CF1 subunits beta and epsilon, respectively, of the chloroplast genome of the brown alga Dictyota dichotoma. Although the coding site of atpE cannot be demonstrated by heterologous Southern hybridizations, a 417 bp reading frame 3' to atpB was identified as the gene atpE by sequence similarities with atpE genes from other sources. A maximum sequence identity of 30% is found between the predicted amino acid sequence of the Dictyota subunit epsilon and the corresponding cyanobacterial subunits. Including conserved amino acid replacements, the Dictyota epsilon subunit exhibits about 70% sequence similarity with the cyanobacterial and land plant subunits. As in cyanobacteria, the atpE gene does not overlap the preceding gene atpB. The deduced amino acid sequence of atpB is 74-79% identical to the corresponding cyanobacterial and chloroplast subunits. Entirely conserved are regions referred to as the catalytic and/or regulatory sites of ATP formation, including interacting regions between subunits alpha and beta. A phylogram predicted from F1/CF1-beta subunits of eleven different organisms suggests a common evolutionary origin of plastids from chlorophytes and brown algae.
We cloned and sequenced the genes of the atpA (atp1) cluster, together with its upstream and downstream sequences, of the chloroplast genome of the cryptomonad Guillardia theta Hill et Wetherbee and used gene signatures, predicted amino acid sequences, and gene arrangements for phylogenetic inferences. The Guillardia atpA cluster contains the genes atpI,H,G,F,D,A in the given order, thus reflecting the cyanobacterial/red algal/chromophyte gene arrangement. In addition, the order of both the upstream sequences rpoB,C1,C2,rps2,tsf and the downstream sequences ycf16/24, which is found exclusively in red algal chloroplast genomes, is conserved in Guillardia. This gene order, which presumably is the result of the reduction process of the cyanobacterial genome following primary endosymbiosis, provides strong evidence for a red algal ancestry of the Guillardia chloroplast and supports the hypothesis of secondary endosymbioses giving rise to chl a+c‐containing algae. The close evolutionary relationship of the chloroplasts of Guillardia and red algae is furthermore supported by the lack of introns, high degrees of sequence similarities, and additional gene signatures, including spacers, gene overlaps, and inverted repeats. Gene cluster analysis, including the ATPase genes together with their upstream and downstream genes, is consistent with a single primary photosynthetic eukaryote that gave rise to all extant algal lineages and land plants by either direct filiation or secondary endosymbioses.
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