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Evolution and gene transfer in purple photosynthetic bacteria.

A concern voiced in connection with recent sequencing of cytochrome c from the Rhodospirillaceae or purple non-sulphur photosynthetic bacteria is that molecular information might be of little use in deciphering bacterial phylogeny because of the possibility of lateral transfer of genes and the consequent scrambling of the genetic record. This could be true for many proteins, of course, but the immediate question is, is it true for cytochroms available for comparison? The evidence suggests that this is probably not so. The disagreement between cytochrome c sequences and the standard taxonomy of the Rhodospirillaceae in Bergey's Manual is not a problem. That reference, as its title indicates, is a manual of determinative rather than evolutionary bacteriology. Its goal is a reproducible system for identification of bacteria. If one claims that these determinative categories also have phylogenetic or evolutionary meaning, this is an assertion that must be proven. The argument outlined below suggests that molecular traits may eventually become more a dependable basis for classification of the Rhodospirillaceae than is gross morphology.

Base Sequence↗

[Changes in blood coagulation in stress and depression--from evolution to gene regulation].

Mental stress and depression are characterized by a hypercoagulable state, which might mediate the increased coronary risk in individuals who feel stressed or depressed. Changes in blood coagulation and fibrinolysis with stress and depression are largely mediated by the sympathetic nervous system via catecholamine and adrenergic receptor activity. Stress might also affect coagulation activity via an influence on the regulation of genes coding for coagulation and fibrinolysis molecules. There is some evidence that non-selective beta-blocking agents decrease hypercoagulability elicited by acute stress. The selective serotonin reuptake inhibitors appear to normalize increased platelet activity in depression. Prospective studies need to show whether there is an association between coagulation abnormalities in stress and depression with hard cardiovascular end-points and whether such an association is favorably affected by therapeutic interventions (e.g., medication, psychotherapy, stress management).

Animals↗

The evolution of genes for pigmentation in African cichlid fishes.

The cichlid fishes in the East African Great Lakes exhibit a great diversity of color patterns, presumably as adaptations to species-specific habitats and/or due to the action of sexual selection on color for species discrimination or female mate choice. To elucidate the mechanisms underlying such pigment pattern diversity, we cloned the cichlid homologs of tyrosinase, endothelin receptor b1, mitf, and Aim1 that previously had been cloned and characterized from pigmentation mutants of zebrafish and medaka fish. Gene sequence analysis among five cichlid species from the Great Lakes shows that the evolutionary rate of amino acid replacement in mitf is the highest of these four genes. We then compared the mitf amino acid replacement rates between species from the lacustrine and tilapiine/steatocranus lineages, and between Lake Malawi- and Victoria-haplochromine cichlids and Lake Tanganyika Lamprologini. We show that the evolutionary rate within the lacustrine lineage is twice that of the tilapiine/steatocranus lineage, but that rates for the Malawi-Victoria haplochromine and Lamprologini lineages are almost the same. These results suggest that the accelerated evolution of mitf might have occurred concomitantly with pigment pattern diversification in Great Lakes species, but not necessarily correlated with species under intense sexual selection on male mating color via female mate choice. Finally, we characterized a novel alternatively spliced variant of cichlid mitf that is similar to a mammalian mitf splice variant generated using alternate splice sites. We suggest that this new variant in cichlids, like that in mammals, encodes an MITF transcriptional factor having higher relative DNA binding affinity. These data provide a novel example of functional convergence in which a particular splice variant is independently generated via alternative splicing of a specific gene in different lineages.

Alternative Splicing↗

Mitochondrial DNA and bindin gene sequence evolution among allopatric species of the sea urchin genus Arbacia.

Sea urchins of the genus Arbacia (order Stirodonta) have discontinuous allopatric distributions ranging over thousands of kilometers. Mitochondrial DNA (mtDNA) sequences were used to reconstruct phylogenetic relationships of four Arbacia species and their geographic populations. There is little evidence of genetic structuring of populations within species, except in two cases at range extremes. The mtDNA sequence differentiation between species suggests that divergence occurred about 4-9 MYA. Gene sequences encoding the sperm protein bindin and its intron were obtained and compared with the mtDNA phylogeny. Sea urchins among the well-studied echinoid order Camarodonta, with degrees of mtDNA divergence similar to those of Arbacia species, are known to have remarkable variation in bindin. However, in Arbacia, little variation in deduced amino acid sequences of bindin was found, indicating that purifying selection acts on the protein. In contrast, bindin intron sequences showed much differentiation, including numerous insertion/deletions. Fertilization experiments performed between a divergent pair of Arbacia species from the Atlantic and Pacific Oceans revealed no evidence of blocks to gamete recognition. In Arbacia, fertilization specificities may have evolved relatively slowly as a result of extensive gene flow within species, greater functional constraint on the bindin polypeptide, or reduced selective pressure for species recognition in singly occurring species.

Amino Acid Sequence↗

Origin and evolution of gene for prolactin-induced protein.

Prolactin-induced protein (PIP) is a small protein secreted into the fluid in several glands. We determined the PIP coding sequences of 5 hominoid species and estimated the numbers of synonymous and nonsynonymous substitutions for each branch of the mammalian PIP gene tree. The branch connecting hominoids and Old World monkeys showed significantly higher nonsynonymous than synonymous substitutions. These changes tended to be accumulated in the fibronectin-binding domain. Many other primate branches also showed higher nonsynonymous than synonymous substitutions, thus suggesting that the PIP genes of primates have experienced some kind of positive selection. We also considered the phylogenetic relationship of the PIP gene with the alpha-2-macroglobulin gene family. The results indicate that the PIP gene arose by partial gene duplication from a member of the alpha-2-macroglobulin gene family after the divergence between amphibians and other tetrapods.

Amino Acid Sequence↗

Expression of the AmphiTcf gene in amphioxus: insights into the evolution of the TCF/LEF gene family during vertebrate evolution.

T-cell factor (TCF) and lymphoid enhancer factors (LEF) genes encode proteins that are transcription factors mediating beta-catenin/Wnt signaling. Whereas mammals have four such genes, the Florida amphioxus (Branchiostoma floridae) apparently has only one such gene (AmphiTcf). From cleavage through early gastrula, cytoplasmic maternal transcripts of this gene are localized toward the animal pole. In gastrulae, AmphiTcf expression begins in the mesendoderm. In neurulae, there is expression in the pharynx, hindgut, anterior notochord, somites, and at the anterior end of the neural plate. In early larvae, expression is detectable in the floor of the diencephalon, notochord, tail bud, forming somites, pharynx, and ciliated pit (a presumed homolog of the vertebrate adenohypophysis). Phylogenetic analysis of TCF/LEF proteins placed AmphiTcf as the sister group of a clade comprising vertebrate Tcf1, Lef1, Tcf3, and Tcf4. Comparison of developmental expression for amphioxus AmphiTcf and vertebrate TCF/LEF genes indicates that this gene family has undergone extensive subfunctionalization and neofunctionalization during vertebrate evolution.

Animals↗

Horizontal gene transfer in microbial genome evolution.

Horizontal gene transfer is the collective name for processes that permit the exchange of DNA among organisms of different species. Only recently has it been recognized as a significant contribution to inter-organismal gene exchange. Traditionally, it was thought that microorganisms evolved clonally, passing genes from mother to daughter cells with little or no exchange of DNA among diverse species. Studies of microbial genomes, however, have shown that genomes contain genes that are closely related to a number of different prokaryotes, sometimes to phylogenetically very distantly related ones. (Doolittle et al., 1990, J. Mol. Evol. 31, 383-388; Karlin et al., 1997, J. Bacteriol. 179, 3899-3913; Karlin et al., 1998, Annu. Rev. Genet. 32, 185-225; Lawrence and Ochman, 1998, Proc. Natl. Acad. Sci. USA 95, 9413-9417; Rivera et al., 1998, Proc. Natl. Acad. Sci. USA 95, 6239-6244; Campbell, 2000, Theor. Popul. Biol. 57 71-77; Doolittle, 2000, Sci. Am. 282, 90-95; Ochman and Jones, 2000, Embo. J. 19, 6637-6643; Boucher et al. 2001, Curr. Opin., Microbiol. 4, 285-289; Wang et al., 2001, Mol. Biol. Evol. 18, 792-800). Whereas prokaryotic and eukaryotic evolution was once reconstructed from a single 16S ribosomal RNA (rRNA) gene, the analysis of complete genomes is beginning to yield a different picture of microbial evolution, one that is wrought with the lateral movement of genes across vast phylogenetic distances. (Lane et al., 1988, Methods Enzymol. 167, 138-144; Lake and Rivera, 1996, Proc. Natl. Acad. Sci. USA 91, 2880-2881; Lake et al., 1999, Science 283, 2027-2028).

Archaea↗

Legumin encoding sequences from the redwood family (Taxodiaceae) reveal precursors lacking the conserved Asn-Gly processing site.

We have cloned and sequenced two different cDNAs encoding legumins from Japanese red cedar (Cryptomeria japonica, Taxodiaceae). The derived amino acid sequences show between 34% and 55% identity when compared with legumins from angiosperms and from Pinaceae, respectively. The predicted precursors are unusual in that they contain potential glycosylation signals, and we have found the corresponding beta-polypeptides actually to be glycosylated. As most outstanding feature one of the precursors is lacking the Asn-Gly processing site which has been assumed to be highly conserved in legumin gene evolution. Legumin encoding sequences amplified from genomic DNA suggest that these unusual precursors are widespread if not ubiquitous in the Taxodiaceae family. From previous reports on legumin precursors with divergent processing sites, on the proteases involved in legumin precursor processing and from the results presented here it is concluded that the Asn-Gly processing site has been acquired rather than conserved during legumin gene evolution.

Amino Acid Sequence↗

Evolution of gene expression patterns in a model of branching morphogenesis.

Branching morphogenesis of the ureteric bud in response to unknown signals from the metanephric mesenchyme gives rise to the urinary collecting system and, via inductive signals from the ureteric bud, to recruitment of nephrons from undifferentiated mesenchyme. An established cell culture model for this process employs cells of ureteric bud origin (UB) cultured in extracellular matrix and stimulated with conditioned media (BSN-CM) from a metanephric mesenchymal cell line (H. Sakurai, E. J. Barros, T. Tsukamoto, J. Barasch, and S. K. Nigam. Proc. Natl. Acad. Sci. USA 94: 6279-6284, 1997.). In the presence of BSN-CM, the UB cells form branching tubular structures reminiscent of the branching ureteric bud. The pattern of gene regulation in this model of branching morphogenesis of the kidney collecting system was investigated using high-density cDNA arrays. Software and analytical methods were developed for the quantification and clustering of genes. With the use of a computational method termed "vector analysis," genes were clustered according to the direction and magnitude of differential expression in n-dimensional log-space. Changes in gene expression in response to the BSN-CM consisted primarily of differential expression of transcription factors with previously described roles in morphogenesis, downregulation of pro-apoptotic genes accompanied by upregulation of anti-apoptotic genes, and upregulation of a small group of secreted products including growth factors, cytokines, and extracellular proteinases. Changes in expression are discussed in the context of a general model for epithelial branching morphogenesis. In addition, the cDNA arrays were used to survey expression of epithelial markers and secreted factors in UB and BSN cells, confirming the largely epithelial character of the former and largely mesenchymal character of the later. Specific morphologies (cellular processes, branching multicellular cords, etc.) were shown to correlate with the expression of different, but overlapping, genomic subsets, suggesting differences in morphogenetic mechanisms at these various steps in the evolution of branching tubules.

Animals↗

Leaf evolution: gases, genes and geochemistry.

AIMS: This Botanical Briefing reviews how the integration of palaeontology, geochemistry and developmental biology is providing a new mechanistic framework for interpreting the 40- to 50-million-year gap between the origination of vascular land plants and the advent of large (megaphyll) leaves, a long-standing puzzle in evolutionary biology. SCOPE: Molecular genetics indicates that the developmental mechanisms required for leaf production in vascular plants were recruited long before the advent of large megaphylls. According to theory, this morphogenetic potential was only realized as the concentration of atmospheric CO2 declined during the late Palaeozoic. Surprisingly, plants effectively policed their own evolution since the decrease in CO2 was brought about as terrestrial floras evolved accelerating the rate of silicate rock weathering and enhancing sedimentary organic carbon burial, both of which are long-term sinks for CO2. CONCLUSIONS: The recognition that plant evolution responds to and influences CO(2) over millions of years reveals the existence of an intricate web of vegetation feedbacks regulating the long-term carbon cycle. Several of these feedbacks destabilized CO2 and climate during the late Palaeozoic but appear to have quickened the pace of terrestrial plant and animal evolution at that time.

Atmosphere↗

Deductions about the number, organization, and evolution of genes in the tomato genome based on analysis of a large expressed sequence tag collection and selective genomic sequencing.

Analysis of a collection of 120,892 single-pass ESTs, derived from 26 different tomato cDNA libraries and reduced to a set of 27,274 unique consensus sequences (unigenes), revealed that 70% of the unigenes have identifiable homologs in the Arabidopsis genome. Genes corresponding to metabolism have remained most conserved between these two genomes, whereas genes encoding transcription factors are among the fastest evolving. The majority of the 10 largest conserved multigene families share similar copy numbers in tomato and Arabidopsis, suggesting that the multiplicity of these families may have occurred before the divergence of these two species. An exception to this multigene conservation was observed for the E8-like protein family, which is associated with fruit ripening and has higher copy number in tomato than in Arabidopsis. Finally, six BAC clones from different parts of the tomato genome were isolated, genetically mapped, sequenced, and annotated. The combined analysis of the EST database and these six sequenced BACs leads to the prediction that the tomato genome encodes approximately 35,000 genes, which are sequestered largely in euchromatic regions corresponding to less than one-quarter of the total DNA in the tomato nucleus.

Arabidopsis↗

Large-scale trends in the evolution of gene structures within 11 animal genomes.

We have used the annotations of six animal genomes (Homo sapiens, Mus musculus, Ciona intestinalis, Drosophila melanogaster, Anopheles gambiae, and Caenorhabditis elegans) together with the sequences of five unannotated Drosophila genomes to survey changes in protein sequence and gene structure over a variety of timescales--from the less than 5 million years since the divergence of D. simulans and D. melanogaster to the more than 500 million years that have elapsed since the Cambrian explosion. To do so, we have developed a new open-source software library called CGL (for "Comparative Genomics Library"). Our results demonstrate that change in intron-exon structure is gradual, clock-like, and largely independent of coding-sequence evolution. This means that genome annotations can be used in new ways to inform, corroborate, and test conclusions drawn from comparative genomics analyses that are based upon protein and nucleotide sequence similarities.

Animals↗

Clustering of tissue-specific genes underlies much of the similarity in rates of protein evolution of linked genes.

Are genes nonrandomly distributed around the genome and might this explain why it was found that, in the mouse genome, proteins of linked genes evolve at similar rates? Anecdotal evidence suggests that the similarity of expression of linked genes might, in part, explain the similarity in their rates of evolution. Immune system genes, for example, are known to evolve at a high rate and sometimes cluster in the genome. Here we develop methods for statistical tests of similarity of expression of linked genes and report that there is a significant tendency for genes of similar expression breadth to be linked. Significantly, when we exclude tissue specific genes from our sample, the similarity in rates of protein evolution of linked genes is greatly diminished, if not abolished. This diminution is not a sampling artifact. In contrast, while half of the immune genes in our sample reside in 1 of 10 immune clusters in the mouse genome, this clustering appears not to affect the extent of local similarity in rates of evolution. The distribution of placentally expressed genes, in contrast, does have an effect.

Animals↗

Functional characterization of nine Norway Spruce TPS genes and evolution of gymnosperm terpene synthases of the TPS-d subfamily.

Constitutive and induced terpenoids are important defense compounds for many plants against potential herbivores and pathogens. In Norway spruce (Picea abies L. Karst), treatment with methyl jasmonate induces complex chemical and biochemical terpenoid defense responses associated with traumatic resin duct development in stems and volatile terpenoid emissions in needles. The cloning of (+)-3-carene synthase was the first step in characterizing this system at the molecular genetic level. Here we report the isolation and functional characterization of nine additional terpene synthase (TPS) cDNAs from Norway spruce. These cDNAs encode four monoterpene synthases, myrcene synthase, (-)-limonene synthase, (-)-alpha/beta-pinene synthase, and (-)-linalool synthase; three sesquiterpene synthases, longifolene synthase, E,E-alpha-farnesene synthase, and E-alpha-bisabolene synthase; and two diterpene synthases, isopimara-7,15-diene synthase and levopimaradiene/abietadiene synthase, each with a unique product profile. To our knowledge, genes encoding isopimara-7,15-diene synthase and longifolene synthase have not been previously described, and this linalool synthase is the first described from a gymnosperm. These functionally diverse TPS account for much of the structural diversity of constitutive and methyl jasmonate-induced terpenoids in foliage, xylem, bark, and volatile emissions from needles of Norway spruce. Phylogenetic analyses based on the inclusion of these TPS into the TPS-d subfamily revealed that functional specialization of conifer TPS occurred before speciation of Pinaceae. Furthermore, based on TPS enclaves created by distinct branching patterns, the TPS-d subfamily is divided into three groups according to sequence similarities and functional assessment. Similarities of TPS evolution in angiosperms and modeling of TPS protein structures are discussed.

Alkyl and Aryl Transferases↗

G+C3 structuring along the genome: a common feature in prokaryotes.

The heterogeneity of gene nucleotide content in prokaryotic genomes is commonly interpreted as the result of three main phenomena: (1) genes undergo different selection pressures both during and after translation (affecting codon and amino acid choice); (2) genes undergo different mutational pressure whether they are on the leading or lagging strand; and (3) genes may have different phylogenetic origins as a result of lateral transfers. However, this view neglects the necessity of organizing genetic information on a chromosome that needs to be replicated and folded, which may add constraints to single gene evolution. As a consequence, genes are potentially subjected to different mutation and selection pressures, depending on their position in the genome. In this paper, we analyze the structuring of different codon usage measures along completely sequenced bacterial genomes. We show that most of them are highly structured, suggesting that genes have different base content, depending on their location on the chromosome. A peculiar pattern of genome structure, with a tendency toward an A+T-enrichment near the replication terminus, is found in most bacterial phyla and may reflect common chromosome constraints. Several species may have lost this pattern, probably because of genome rearrangements or integration of foreign DNA. We show that in several species, this enrichment is associated with an increase of evolutionary rate and we discuss the evolutionary implications of these results. We argue that structural constraints acting on the circular chromosome are not negligible and that this natural structuring of bacterial genomes may be a cause of overestimation in lateral gene transfer predictions using codon composition indices.

Base Composition↗

Molecular genetic aspects of tetraploidy in the common carp Cyprinus carpio.

Genome duplications are believed to have occurred on multiple occasions in vertebrate evolution. Studies of duplicate gene loci in tetraploid animals may reveal important general aspects of gene duplication, an important mode of gene evolution in metazoans. The common carp Cyprinus carpio has twice as many chromosomes as most other cyprinid fishes due to tetraploidization previously estimated to have occurred 50 Myr ago. Our sequence analyses of duplicate carp loci suggest that the tetraploidization took place less than 16 Myr ago. This is further supported by sequence comparisons with the diploid grass carp, which seems to have diverged from the common carp approximately 19 Myr ago. Duplicate loci appear to remain expressed for millions of years and may accumulate mutations leading to drastic amino acid replacements as shown here for somatotropin. Therefore, both loci should always be characterized in molecular studies of tetraploid animals such as goldfish, salmonid fishes, and Xenopus laevis. The long life of duplicate genes may explain the occurrence of numerous large multigene families in higher metazoans.

Actins↗

Convergent evolution. The gene structure of Sulculus 41 kDa myoglobin is homologous with that of human indoleamine dioxygenase.

The abalone Sulculus diversicolor contains abundant myoglobin in its buccal mass. The myoglobin consists of 377 amino acid residues and has a molecular mass of 41 000 Da, 2.5 times larger than that of other myoglobins. Sulculus myoglobin can bind oxygen reversibly, and the P50 was determined to be 3.8 mmHg at 20 degrees C and pH 7.4, showing that the oxygen affinity of Sulculus myoglobin is lower than those of vertebrate and invertebrate myoglobins. The cDNA-derived amino acid sequence showed no significant homology with those of any other invertebrate myoglobins and hemoglobins, but surprisingly showed 35% homology with a vertebrate tryptophan-degrading enzyme, indoleamine dioxygenase (IDO). The structure of the Sulculus myoglobin gene has been determined to consist of 14 exons and 13 introns (15.3 kbp). Compared with the gene of human IDO (10 exon-9 intron structure), the splice junctions of 7 introns were exactly conserved between the two genes, suggesting that these introns have been conserved for at least 600 million years. The Sulculus gene has 5 additional introns, one of which is located outside the coding region. From these results we conclude that Sulculus myoglobin evolved from an IDO gene and represents a typical case of functional convergence. Comparison of the amino acid sequence of each exon of Sulculus myoglobin with those of usual globin sequences showed that there is no significant evolutionary relationship between them. The IDO-like myoglobin is unexpectedly widely distributed among gastropodic molluscs, such as Sulculus, Nordotis, Battilus, Omphalius and Chlorostoma.

Amino Acid Sequence↗

Unifying measures of gene function and evolution.

Recent genome analyses revealed intriguing correlations between variables characterizing the functioning of a gene, such as expression level (EL), connectivity of genetic and protein-protein interaction networks, and knockout effect, and variables describing gene evolution, such as sequence evolution rate (ER) and propensity for gene loss. Typically, variables within each of these classes are positively correlated, e.g. products of highly expressed genes also have a propensity to be involved in many protein-protein interactions, whereas variables between classes are negatively correlated, e.g. highly expressed genes, on average, evolve slower than weakly expressed genes. Here, we describe principal component (PC) analysis of seven genome-related variables and propose biological interpretations for the first three PCs. The first PC reflects a gene's 'importance', or the 'status' of a gene in the genomic community, with positive contributions from knockout lethality, EL, number of protein-protein interaction partners and the number of paralogues, and negative contributions from sequence ER and gene loss propensity. The next two PCs define a plane that seems to reflect the functional and evolutionary plasticity of a gene. Specifically, PC2 can be interpreted as a gene's 'adaptability' whereby genes with high adaptability readily duplicate, have many genetic interaction partners and tend to be non-essential. PC3 also might reflect the role of a gene in organismal adaptation albeit with a negative rather than a positive contribution of genetic interactions; we provisionally designate this PC 'reactivity'. The interpretation of PC2 and PC3 as measures of a gene's plasticity is compatible with the observation that genes with high values of these PCs tend to be expressed in a condition- or tissue-specific manner. Functional classes of genes substantially vary in status, adaptability and reactivity, with the highest status characteristic of the translation system and cytoskeletal proteins, highest adaptability seen in cellular processes and signalling genes, and top reactivity characteristic of metabolic enzymes.

Animals↗