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HLA-A*8001 is a member of a newly discovered ancient family of HLA-A alleles.

A member of a new HLA-A locus family, HLA-A*8001, has been characterized from 3 African-American individuals expressing a unique HLA-A serologic specificity. The HLA-A*8001 sequence is most closely related to alleles of the HLA-A1/3/11 family, although it also contains residues characteristic of the HLA-A2/28, -A9, -A10, and -A19 families. More importantly, the HLA-A*8001 sequence contains four unique nonsynonymous (amino acid replacing) nucleotide substitutions absent from all other primate A locus alleles. In addition, five other nucleotide substitutions, four nonsynonymous and one synonymous (silent), observed only in non-human primate A locus alleles were found. Neighbor-joining tree analysis of HLA-A*8001 supports the notion that the HLA-A*8001 allele is a member of a new HLA-A locus family which was derived from the ancestral A3 lineage but diverged early in the evolution of the HLA-A locus.

Alleles↗

Complex evolution of photosynthesis.

The origin of photosynthesis is a fundamental biological question that has eluded researchers for decades. The complexity of the origin and evolution of photosynthesis is a result of multiple photosynthetic components having independent evolutionary pathways. Indeed, evolutionary scenarios have been established for only a few photosynthetic components. Phylogenetic analysis of Mg-tetrapyrrole biosynthesis genes indicates that most anoxygenic photosynthetic organisms are ancestral to oxygen-evolving cyanobacteria and that the purple bacterial lineage may contain the most ancestral form of this pigment biosynthesis pathway. The evolutionary path of type I and type II reaction center apoproteins is still unresolved owing to the fact that a unified evolutionary tree cannot be generated for these divergent reaction center subunits. However, evidence for a cytochrome b origin for the type II reaction center apoproteins is emerging. Based on the combined information for both photopigments and reaction centers, a unified theory for the evolution of reaction center holoproteins is provided. Further insight into the evolution of photosynthesis will have to rely on additional broader sampling of photosynthesis genes from divergent photosynthetic bacteria.

Amino Acid Sequence↗

Do early branching lineages signify ancestral traits?

A reverence for ancestors that has pre-occupied humans since time immemorial persists to the present. Reconstructing ancestry is the focus of many biological studies but failure to distinguish between present-day descendants and long-dead ancestors has led to incorrect interpretation of phylogenetic trees. This has resulted in erroneous reconstruction of traits such as morphology and ancestral areas. Misinterpretation becomes evident when authors use the terms 'basal' or 'early diverging' to refer to extant taxa. Here, we discuss the correct interpretation of trees and methods for reconstructing the ancestral features of organisms using recently developed statistical models. These models can be inaccurate unless they use information that is independent of phylogenies, such as genetics, molecular and developmental biology, functional morphology, geological and climatic processes, and the fossil record.

Journal Article↗

Radiation and diversification within the Ligularia-Cremanthodium-Parasenecio complex (Asteraceae) triggered by uplift of the Qinghai-Tibetan Plateau.

The Ligularia-Cremanthodium-Parasenecio (L-C-P) complex of the Tussilagininae (Asteraceae: Senecioneae) contains more than 200 species that are endemic to the Qinghai-Tibetan Plateau in eastern Asia. These species are morphologically distinct; however, their relationships appear complex. A phylogenetic analysis of members of the complex and selected taxa of the tribe Senecioneae was conducted using chloroplast (ndhF and trnL-F) and nuclear (ITS) sequences. Phylogenetic trees were constructed from individual and combined datasets of the three different sequences. All analyses suggested that Doronicum, a genus that has been included in the Tussilagininae, should be excluded from this subtribe and placed at the base of the tribe Senecioneae. In addition, the Tussilagininae should be broadly circumscribed to include the Tephroseridinae. Within the expanded Tussilagininae containing all 13 genera occurring in eastern Asia, Tussilago and Petasites diverged early as a separate lineage, while the remaining 11 genera comprise an expanded L-C-P complex clade. We suggest that the L-C-P clade, which is largely unresolved, most likely originated as a consequence of an explosive radiation. The few monophyletic subclades identified in the L-C-P clade with robust support further suggest that some genera of Tussilagininae from eastern Asia require generic re-circumscriptions given the occurrence of subclades containing species of the same genus in different parts of the phylogentic tree due to homoplasy of important morphological characters used to delimit them. Molecular-clock analyses suggest that the explosive radiation of the L-C-P complex occurred mostly within the last 20 million years, which falls well within the period of recent major uplifts of the Qinghai-Tibetan Plateau between the early Miocene to the Pleistocene. It is proposed that significant increases in geological and ecological diversity that accompanied such uplifting, most likely promoted rapid and continuous allopatric speciation in small and isolated populations, and allowed fixation or acquisition of similar morphological characters within unrelated lineages. This phenomenon, possibly combined with interspecific diploid hybridization because of secondary sympatry during relatively stable stages between different uplifts, could be a major cause of high species diversity in the Qinghai-Tibetan Plateau and adjacent areas of eastern Asia.

Asteraceae↗

Three globin lineages belonging to two structural classes in genomes from the three kingdoms of life.

Although most globins, including the N-terminal domains within chimeric proteins such as flavohemoglobins and globin-coupled sensors, exhibit a 3/3 helical sandwich structure, many bacterial, plant, and ciliate globins have a 2/2 helical sandwich structure. We carried out a comprehensive survey of globins in the genomes from the three kingdoms of life. Bayesian phylogenetic trees based on manually aligned sequences indicate the possibility of past horizontal globin gene transfers from bacteria to eukaryotes. blastp searches revealed the presence of 3/3 single-domain globins related to the globin domains of the bacterial and fungal flavohemoglobins in many bacteria, a red alga, and a diatom. Iterated psi-blast searches based on groups of globin sequences found that only the single-domain globins and flavohemoglobins recognize the eukaryote 3/3 globins, including vertebrate neuroglobins, alpha- and beta-globins, and cytoglobins. The 2/2 globins recognize the flavohemoglobins, as do the globin coupled sensors and the closely related single-domain protoglobins. However, the 2/2 globins and the globin-coupled sensors do not recognize each other. Thus, all globins appear to be distributed among three lineages: (i) the 3/3 plant and metazoan globins, single-domain globins, and flavohemoglobins; (ii) the bacterial 3/3 globin-coupled sensors and protoglobins; and (iii) the bacterial, plant, and ciliate 2/2 globins. The three lineages may have evolved from an ancestral 3/3 or 2/2 globin. Furthermore, it appears likely that the predominant functions of globins are enzymatic and that oxygen transport is a specialized development that accompanied the evolution of metazoans.

Bayes Theorem↗

Fixation times of retroposons in the ribosomal DNA spacer of human and other primates.

We have investigated the presence/absence of two types of retroposed sequences found in human ribosomal DNA in equivalent positions in chimpanzee, gorilla, orangutan, gibbon, and rhesus monkey rDNA. These sequences are one pseudogene derived from the single-copy cdc27hs gene and seven complete Alu elements. The 2-kb pseudogene is present in the apes but not in Old World monkeys, indicating fixation in an ape ancestor. Five of the Alu elements are shared by the whole set of primates studied, indicating insertion and fixation prior to the split of the ape and Old World monkey lineages. One is absent only from the rhesus monkey rDNA, and another is absent from both gibbon and rhesus rDNA, indicating fixation at different times in primate evolutionary history. Since branching times for the primate phylogenetic tree are known from a combination of the fossil record and multiple molecular data sets, it is possible to compare Alu fixation times determined from the phylogenetic information with those calculated from Alu element mutation rates.

Animals↗

Phylogeny and evolution of the Sulidae (Aves:Pelecaniformes): a test of alternative modes of speciation.

Although the allopatric model of speciation is widely accepted, it does not provide a satisfactory explanation for many evolutionary phenomena. Several alternative models exist, but they remain largely untested for vertebrate animals. In the present paper, a molecular phylogeny was used to test competing models of speciation in a seabird family, the Sulidae. A segment including 807 base pairs of the mitochondrial cytochrome b gene was sequenced from all extant sulid species, and phylogenetic methods were used to test model-specific predictions regarding tree topologies, distributions of sister taxa, timing of vicariant events, and comparative biology. Both the neighbor-joining and parsimony analyses placed sequences of gannets (Morus spp.) and boobies of the genus Sula in separate, monophyletic lineages. Sequences of Cape (M. capensis) and Australasian (M. serrator) gannets clustered together, and the sequence of Abbott's booby (Papasula abbotti) was basal to those of the gannets. Sequences of blue-footed (S. nebouxii) and Peruvian (S. variegata) boobies were sisters and formed a monophyletic group with the masked booby (S. dactylatra). The red-footed booby (S. sula) sequence was the most divergent of the Sula boobies. All relationships received strong support from standard-error tests and bootstrap analysis. Substitution rates were similar to those suggested for mammals and suggested that most lineages arose within the last 3 million years. Lineage divergence events for which the mode of speciation could be deduced did not fit the predictions of either allopatric or sympatric models, but apparently involved either peripatric or parapatric processes.

Animals↗

The evolutionary relationships among known life forms.

Sequences of small subunit (SSU) and large subunit (LSU) ribosomal RNA genes from archaebacteria, eubacteria, and the nucleus, chloroplasts, and mitochondria of eukaryotes have been compared in order to identify the most conservative positions. Aligned sets of these positions for both SSU and LSU rRNA have been used to generate tree diagrams relating the source organisms/organelles. Branching patterns were evaluated using the statistical bootstrapping technique. The resulting SSU and LSU trees are remarkably congruent and show a high degree of similarity with those based on alternative data sets and/or generated by different techniques. In addition to providing insights into the evolution of prokaryotic and eukaryotic (nuclear) lineages, the analysis reported here provides, for the first time, an extensive phylogeny of the mitochondrial lineage.

Base Sequence↗

Molecular phylogeny of the free-living archezoan Trepomonas agilis and the nature of the first eukaryote.

We have sequenced the small ribosomal subunit RNA gene of the diplozoan Trepomonas agilis. This provides the first molecular information on a free-living archezoan. We have performed a phylogenetic analysis by maximum likelihood, parsimony, and distance methods for all available nearly complete archezoan small subunit ribosomal RNA genes and for representatives of all major groups of more advanced eukaryotes (metakaryotes). These show Diplozoa as the earliest-diverging eukaryotic lineage, closely followed by microsporidia. Trepomonas proves to be much more closely related to Hexamita, and, to a lesser degree, to Spironucleus, than to Giardia. The close relationship between the free-living Trepomonas on our trees and the parasites Hexamita inflata and Spironucleus refutes the idea that the early divergence of the amitochondrial Archezoa is an artefact caused by parasitism. The deep molecular divergence between the three phagotrophic genera with two cytostomes (Hexamita, Trepomonas, Spironucleus) and the saprotrophic Giardia that lacks cytostomes is in keeping with the classical evidence for a fundamental difference in the symmetry of the cytoskeleton between the two groups. We accordingly separate the two groups as two orders: Distomatida for those with two cytostomes/cytopharynxes and Giardiida ord. nov. for Giardia and Octomitus that lack these, and divide each order into two families. We suggest that this fundamental divergence in manner of feeding and in the symmetry of the cytoskeleton evolved in a free-living diplozoan very early indeed in the evolution of the eukaryotic cell, possibly very soon after the origin of the diplokaryotic state (having two nuclei linked together firmly by the cytoskeleton) and before the evolution of parasitism by distomatids and giardiids, which may have colonized animal guts independently. We discuss the possible relationship between the two archezoan phyla (Metamonada and Microsporidia) and the nature of the first eukaryotic cell in the light of our results and other recent molecular data.

Animals↗

Relaxed molecular clocks for dating historical plant dispersal events.

Age estimation from molecular sequences has emerged as a powerful tool for inferring when a plant lineage arrived in a particular area. Knowing the tenure of lineages within a region is key to understanding the evolution of traits, the evolution of biotic interactions, and the evolution of floras. New analytical methods model change in substitution rates along individual branches of a phylogenetic tree by combining molecular data with time constraints, usually from fossils. These "relaxed clock" approaches can be applied to several gene regions that need not all have the same substitution rates, and they can also incorporate multiple simultaneous fossil calibrations. Since 1995, at least 100 plant biogeographic studies have used molecular-clock dating, and about a fifth has used relaxed clocks. Many of these report evidence of long-distance dispersal. Meta-analyses of studies from the same geographic region can identify directional biases because of prevailing wind or water currents and the relative position and size of landmasses.

Geography↗

Multiple origins of the ascidian-Prochloron symbiosis: molecular phylogeny of photosymbiotic and non-symbiotic colonial ascidians inferred from 18S rDNA sequences.

In the tropics, certain didemnid ascidians harbor the prokaryotic photosymbiont Prochloron. To date, this photosymbiosis has been found in four didemnid genera that include non-symbiotic species. Here, we report the molecular phylogeny of symbiotic and non-symbiotic didemnids based on their 18S rDNA sequences. The data cover all four genera containing symbiotic species and one other genus comprised of only non-symbiotic species. Near-complete nucleotide sequences of 18S rDNAs were determined for four non-didemnid species and 52 didemnid samples (five genera), including 48 photosymbiotic samples collected from the Ryukyu Archipelago, the Great Barrier Reef, Hawaii, and Bali. Our phylogenetic trees indicated a monophyletic origin of the family Didemnidae, as well as each of the didemnid genera. The results strongly support the hypothesis that establishment of the ascidian-Prochloron symbiosis occurred independently in the Didemnidae lineage at least once in each of the genera that possess symbiotic species.

Animals↗

Fungus-specific microsatellite primers of lichens: application for the assessment of genetic variation on different spatial scales in Lobaria pulmonaria.

We isolated 12 microsatellite loci for the epiphytic lichen-forming ascomycete Lobaria pulmonaria and studied their patterns of variation within and among populations from Canada and Switzerland. Even though several microsatellites exhibited high levels of variability at different spatial scales, we did not find any evidence for intrathalline variation. Most of the genetic variation was attributed to differences among individuals within populations. High genetic variation was also detected among L. pulmonaria samples taken from individual trees, suggesting that either multiple colonization events had occurred or that local recombination is frequent. The geographically structured distribution of alleles from several microsatellites indicated that L. pulmonaria from Canada and Switzerland represent two distinct evolutionary lineages. The potential to identify multiple alleles, and their transferability to closely related species, make microsatellites an ideal tool to study dispersal, population differentiation, and microevolution in lichens.

Alleles↗

Mitochondrial DNA sequences of five squamates: phylogenetic affiliation of snakes.

Complete or nearly complete mitochondrial DNA sequences were determined from four lizards (Western fence lizard, Warren's spinytail lizard, Terrestrial arboreal alligator lizard, and Chinese crocodile lizard) and a snake (Texas blind snake). These genomes had a typical gene organization found in those of most mammals and fishes, except for a translocation of the glutamine tRNA gene in the blind snake and a tandem duplication of the threonine and proline tRNA genes in the spinytail lizard. Although previous work showed the existence of duplicate control regions in mitochondrial DNAs of several snakes, the blind snake did not have this characteristic. Phylogenetic analyses based on different tree-building methods consistently supported that the blind snake and a colubrid snake (akamata) make a sister clade relative to all the lizard taxa from six different families. An alternative hypothesis that snakes evolved from a lineage of varanoids was not favored and nearly statistically rejected by the Kishino-Hasegawa test. It is therefore likely that the apparent similarity of the tongue structure between snakes and varanoids independently evolved and that the duplication of the control region occurred on a snake lineage after divergence of the blind snake.

Animals↗

Evidence for directional selection acting on pheromone-binding proteins in the genus Choristoneura.

Patterns of nucleotide variation consistent with the action of natural selection have been discovered at a number of different gene loci. Here, pheromone-binding proteins (PBPs) are examined to determine if selection has acted to fix amino acid changes in PBPs in lineages in which pheromone changes have occurred. PBPs from five different species of moths in the genus Choristoneura were sequenced, along with the PBP of Argyrotaenia velutinana, which serves as an outgroup. Three independent major pheromone changes are represented within this group of five Choristoneura species. Two different lineages show evidence for selection based on polymorphism and divergence comparisons and comparisons of rates of replacement evolution to silent and noncoding evolution. Along one of these lineages, leading to Choristoneura fumiferana, there has been a change to an aldehyde pheromone from an acetate pheromone. The second branch does not appear to be associated with a major pheromone change. Other branches in the tree show a trend toward greater replacement fixation than expected under neutrality. This trend could reflect undetected selective events within this group of PBPs. Selection appears to have acted to fix amino acid changes in the PBP of moths from the genus Choristoneura, but it is not clear that this selection is due to pheromone changes between species.

Amino Acid Sequence↗

A covarion-based method for detecting molecular adaptation: application to the evolution of primate mitochondrial genomes.

A new method for detecting site-specific variation of evolutionary rate (the so-called covarion process) from protein sequence data is proposed. It involves comparing the maximum-likelihood estimates of the replacement rate of an amino acid site in distinct subtrees of a large tree. This approach allows detection of covarion at the gene or the amino acid levels. The method is applied to mammalian-mitochondrial-protein sequences. Significant covarion-like evolution is found in the (simian) primate lineage: some amino acid positions are fast-evolving (i.e. unconstrained) in non-primate mammals but slow-evolving (i.e. highly constrained) in primates, and some show the opposite pattern. Our results indicate that the mitochondrial genome of primates reached a new peak of the adaptive landscape through positive selection.

Animals↗

Specialized cheating of the ectomycorrhizal symbiosis by an epiparasitic liverwort.

Many non-photosynthetic vascular plants in 10 diverse families obtain all of their carbon from fungi, but in most cases the fungi and the ultimate sources of carbon are unknown. In a few cases, such plants have been shown to be epiparasitic because they obtain carbon from neighbouring green plants through shared mycorrhizal fungi. In all such cases, the epiparasitic plants have been found to specialize upon narrow lineages of ecto- or arbuscular mycorrhizal fungi. Here we show that a non-vascular plant, the non-photosynthetic liverwort Cryptothallus mirabilis, is epiparasitic and is specialized on Tulasnella species that form ectomycorrhizae with surrounding trees at four locations in England, France and Portugal. By using microcosm experiments we show that the interaction with Tulasnella is necessary for growth of Cryptothallus, and by using labelling experiments we show that (14)CO(2) provided to birch seedlings is transferred to Cryptothallus by Tulasnella. This is one of the first documented cases of epiparasitism by a non-vascular plant and of ectomycorrhizal formation by Tulasnella. These results broaden the emerging association between epiparasitism and mycorrhizal specialization into a new class of plants and a new order of fungi.

Carbon↗

Molecular evidence for novel planctomycete diversity in a municipal wastewater treatment plant.

We examined anoxic and aerobic basins and an anaerobic digestor of a municipal wastewater treatment plant for the presence of novel planctomycete-like diversity. Three 16S rRNA gene libraries were constructed by using a 16S rRNA-targeted universal reverse primer and a forward PCR primer specific for Planctomyces: Phylogenetic analysis of 234 16S rRNA gene sequences defined 110 operational taxonomic units. The majority of these sequences clustered with the four known genera, Pirellula (32%), Planctomyces (18.4%), Gemmata (3.8%), and Isosphaera (0.4%). More interestingly, 42.3% of the sequences appeared to define two distantly separated monophyletic groups. The first group, represented by 35.5% of the sequences, was related to the Planctomyces group and branched as a monophyletic cluster. It exhibited between 11.9 and 20.3% 16S rRNA gene sequence dissimilarity in comparisons with cultivated planctomycetes. The second group, represented by 6.8% of the sequences, was deeply rooted within the Planctomycetales tree. It was distantly related to the anammox sequences (level of dissimilarity, 20.3 to 24.4%) and was a monophyletic cluster. The retrieved sequences extended the intralineage phylogenetic depth of the Plantomycetales from 23 to 30.6%. The lineages described here may have a broad diversity of undiscovered biochemical and metabolic novelty. We developed a new 16S rRNA-targeted oligonucleotide probe and localized members of one of the phylogenetic groups using the fluorescent in situ hybridization technique. Our results indicate that activated sludge contains very diverse representatives of this group, which grow under aerobic and anoxic conditions and even under anaerobic conditions. The majority of species in this group remain poorly characterized.

Bacteria↗

Plaque morphology of Italian H7N1 LPAI isolates in MDCK cells and in primary cells of different avian species.

The morphology of plaques induced by Italian, H7N1, low-pathogenic avian influenza (LPAI) viruses belonging to different lineages was investigated in primary chicken, turkey, Muscovy duck, and mallard duck kidney cells and in MDCK cells in the absence of trypsin. LPAI isolates were selected on the basis of the location in the phylogenetic tree: 977/V99 (located at the root, no additional glycosylation site [nAGS]), 2379N99 (AGS in position 123), and 3675/V99 (AGS in position 149). Different isolates did not induce plaques with a statistically significant different size in MDCK cells. However, in primary cells of different avian origin, the presence or absence of AGS significantly influenced plaque size. Generally speaking, 977N99 was the least efficient at plaquing in all cells, while 2379/V99 (AGS in position 123) plaqued more efficiently in turkey cells and 3675/V99 (AGS in position 149) in chicken cells. The presence of either AGS induced statistically significant larger plaques in cells of waterfowl origin.

Animals↗