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D Graur

Publications and source records attributed to D Graur.

At least 19 recordsLinked to original sources

Constructing phylogenies from quartets: elucidation of eutherian superordinal relationships.

In this work we present two new approaches for constructing phylogenetic trees. The input is a list of weighted quartets over n taxa. Each quartet is a subtree on four taxa, and its weight represents a confidence level for the specific topology. The goal is to construct a binary tree with n leaves such that the total weight of the satisfied quartets is maximized (an NP hard problem). The first approach we present is based on geometric ideas. Using semidefinite programming, we embed the n points on the n-dimensional unit sphere, while maximizing an objective function. This function depends on Euclidean distances between the four points and reflects the quartet topology. Given the embedding, we construct a binary tree by performing geometric clustering. This process is similar to the traditional neighbor joining, with the difference that the update phase retains geometric meaning: When two neighbors are joined together, their common ancestor is taken to be the center of mass of the original points. The geometric algorithm runs in poly(n) time, but there are no guarantees on the quality of its output. In contrast, our second algorithm is based on dynamic programming, and it is guaranteed to find the optimal tree (with respect to the given quartets). Its running time is a modest exponential, so it can be implemented for modest values of n. We have implemented both algorithms and ran them on real data for n = 15 taxa (14 mammalian orders and an outgroup taxon). The two resulting trees improve previously published trees and seem to be of biological relevance. On this dataset, the geometric algorithm produced a tree whose score is 98.2% of the optimal value on this input set (72.1% vs. 73.4%). This gives rise to the hope that the geometric approach will prove viable even for larger cases where the exponential, dynamic programming approach is no longer feasible.

Algorithms

Patterns and rates of indel evolution in processed pseudogenes from humans and murids.

Patterns and rates of indel (deletions and insertions) evolution were characterized in 156 independently derived processed pseudogenes from humans and murids (mice and rats). A total of 441 deletions and 161 insertions were unambiguously identified. On a subset of 109 pseudogenes, we verified and confirmed the assumption that indels occur almost exclusively in the pseudogene and, therefore, in comparisons between pseudogenes and their functional paralogs, it is possible to assign polarity to the indel event. By comparing the characteristics of terminal truncations with those of internal deletions, we find support for the hypothesis that truncations are generated through a different pathway than internal deletions. The number of deletions and insertions per pseudogene was found to increase monotonically with time. Deletions occur on average once every 40 nucleotide substitutions, whereas insertions are much rarer, occurring once every 100 substitutions, indicating that the mechanisms involved in deletion formation are most probably different from those responsible for the formation of insertions. The age of the pseudogene, however, explained only 20 and 13%, respectively, of the variation in the number of deletions and insertions per site, indicating that factors other than evolutionary time may play a significant role in the evolutionary dynamics of indel accumulation. Since the rate of substitution has been previously shown to be higher in murids than in humans, we deduce that deletions and insertions accumulate proportionally faster in murids than in humans. Deletions and insertions in murid and human genomes do not contribute significantly to genome size.

Animals

Evolutionary affinities of the order Perissodactyla and the phylogenetic status of the superordinal taxa Ungulata and Altungulata.

Contrary to morphological claims, molecular data indicate that the order Perissodactyla (e.g., horses, rhinoceroses, and tapirs) is neither part of the superordinal taxon Paenungulata (Sirenia, Proboscidea, and Hyracoidea) nor an immediate outgroup of the paenungulates. Rather, Perissodactyla is closer to Carnivora and Cetartiodactyla (Cetacea+Artiodactyla) than it is to the paenungulates. Therefore, two morphologically defined superordinal taxa, Altungulata (Proboscidea, Sirenia, Hyracoidea, and Perissodactyla) and Ungulata (Altungulata and Cetartiodactyla), are invalidated. Perissodactyla, Carnivora, and Cetartiodactyla are shown to constitute a rather tight trichotomy. However, a molecular analysis of 36 protein sequences with a total concatenated length of 7885 aligned amino acids indicates that Perissodactyla is closer to Cetartiodactyla than either taxa is to Carnivora. The relationships among Paenungulata, Primates, and the clade consisting of Perissodactyla, Carnivora, and Cetartiodactylaa could not be resolved on the basis of the available data.

Amino Acid Sequence

Phylogenetic position of the order Lagomorpha (rabbits, hares and allies)

Ever since they have been classified as ruminants in the Old Testament (Leviticus 11:6, Deuteronomy 14:7) and equated with hyraxes in the vulgate Latin translation, rabbits and their relatives (order Lagomorpha) have frequently experienced radical changes in taxonomic rank. By using 91 orthologous protein sequences, we have attempted to answer the classical question "What, if anything, is a rabbit?". Here we show that Lagomorpha is significantly more closely related to Primates and Scandentia (tree shrews) than it is to rodents. This newly determined phylogenetic position invalidates the superordinal taxon Glires (Lagomorpha + Rodentia), and indicates that the morphological 'synapomorphies' previously used to cluster rodents and lagomorphs into Glires, may actually represent symplesiomorphies or homoplasies that are of no phylogenetic value. This raises the possibility that the ancestral eutherian morphotype may have possessed many rodent-like morphological characters.

Animals

The "Phoca standard": an external molecular reference for calibrating recent evolutionary divergences.

Comparison of the complete mitochondrial DNA (mtDNA) of the high-Arctic ringed seal (Phoca hispida) and the sub-Arctic harbour (P. vitulina) and grey (Halichoerus grypus) seals shows that they are genetically equidistant from one another. We relate the evolutionary divergence of the three species to expanding glaciation in the Arctic Basin and establish, in conjunction with mtDNA data, a standard reference for calibration of recent divergence events among mammalian taxa. In the present study, we apply the "Phoca standard" to the dating of divergences within the hominid phylogenetic tree. After determining the relative rates of substitution over all mitochondrial protein-coding genes in the different evolutionary lineages, we estimate that humans and chimpanzees diverged from each other 6.1 Mya (95% confidence limits: 5.2-6.9 Mya). The corresponding lower-limit divergence between common chimpanzee, Pan troglodytes, and pygmy chimpanzee, P. paniscus, occurred 3 (2.4-3.6) Mya, and the primary split within the P. troglodytes complex 1.6 (1.3-2.0) Mya. The analyses suggest that the split between Gorilla and Pan/Homo occurred 8.4 (7.3-9.4) Mya. They also suggest that Pongo (orangutan) and the lineage leading to gorillas, chimpanzees, and humans diverged 18.1 (16.5-19.6) Mya. The present analysis is independent of the hominid paleontological record and inferential morphological interpretations and thus is a novel approach to the lower-limit dating of recent divergences.

Animals

Unbiased estimation of symmetrical directional mutation pressure from protein-coding DNA.

The most generally applicable procedure for obtaining estimates of the symmetrical, or strandnonspecific, directional mutation pressure (microD) on protein-coding DNA sequences is to determine the G+C content at synonymous codon sites (Psyn), and to divide Psyn by twice the arithmetic mean of the G+C content at synonymous codon sites of a large number of randomly generated, synonymously coding DNA sequences (Psyn). Unfortunately, the original procedure yields biased estimates of Psyn and microD and is computationally expensive. We here present a fast procedure for estimating unbiased microD values. The procedure employs direct calculation of Psyn (approximately Psyn) and two normalization procedures, one for Psyn < or = Psyn and another for Psyn > or = Psyn. The normalization removes a bias sometimes caused by codons specifying arginine, asparagine, isoleucine, and leucine. Consequently, comparison of protein-coding genes that are translated using different genetic codes is facilitated.

Animals

Mitochondrial-DNA sequence evidence on the phylogeny of Australian jack-jumper ants of the Myrmecia pilosula complex.

Australian ants of the Myrmecia pilosula species complex include some individuals (in M. croslandi) with the lowest possible metazoan chromosome number of 2n = 2. Others in this cluster of sibling species have much higher numbers, the known maximum being 2n = 32. Two species (M. pilosula and M. 'banksi') are believed on cytogenetic and morphological grounds to have hybridized over a long period. To investigate the phylogeny and age of this group relative to the congeneric outgroup species M. gulosa, we sequenced part of the cytochrome b gene and the intergenic sequence between it and a primer anchored on the nearby tRNA(UCNSer) gene and analyzed the coding region using bootstrapped parsimony and neighbor-joining trees using the numbers of synonymous and nonsynonymous codons per site. The intergenic space demonstrated a profusion of repeated sequences, and only very closely related sequences (as judged by that for cytochrome b) showed detectable similarity at this almost 100% A+T region. In agreement with predictions from karyotype studies, the phylogenetic analyses showed that M. croslandi is the sister group to the other siblings; the time of separation of M. croslandi from the rest of the pilosula group is unexpectedly ancient. Other relationships were poorly resolved, but the results suggest that M. 'banski' and M. pilosula cluster together, as expected on cytogenetic grounds, and the tentative suggestion of close affinity of the M. pilosula samples and two "PB" samples supports derivation of PB from female M. pilosula and male M. 'banksi.'

Animals

Multiple cDNAs of wheat voltage-dependent anion channels (VDAC): isolation, differential expression, mapping and evolution.

The mitochondrial outer membrane of eukaryotic cells contains voltage-dependent anion channels (VDAC) also termed porins. Three cDNAs from wheat (Triticum aestivum) were isolated and sequenced (Tavdac 1-3). They share 65% similarity of their amino acid sequences, and therefore they probably represent isoforms. The deduced amino acid sequence of one of the cDNAs was found to be identical to the purified VDAC protein from wheat mitochondria [8]. Secondary structure analysis of the deduced amino acid sequences of the three vdac cDNAs revealed a characteristic alpha helix at their N-terminal and beta-barrel cylinders characteristic of VDAC channels. The Tavdac cDNAs are differentially expressed in meristematic tissues. The transcript levels of Tavdac 1 in all wheat tissues is at least 2.5-fold higher than Tavdac 2 and Tavdac 3. Tavdac 2 has a low level of expression in all floral tissues whereas Tavdac 3 is highly expressed in anthers. This is the first report on differential expression of vdac genes in plants. The Tavdac genes have been mapped on the wheat genome. Tavdac 1 is located on the long arm of chromosome 5, Tavdac 2 on the long arm of chromosome 1 and Tavdac 3 on the long arm of chromosome 3. A phylogenetic reconstruction indicates that vdac genes underwent numerous duplication events throughout their evolution. All duplications occurred after the separation of plants from animals and fungi, and no orthologous genes are shared among phyla. Within plants, some of the vdac gene duplications probably occurred before the monocotydelon-dicotydelon split.

Amino Acid Sequence

Analysis of directional mutation pressure and nucleotide content in mitochondrial cytochrome b genes.

We present a new approach for analyzing directional mutation pressure and nucleotide content in protein-coding genes. Directional mutation pressure, the heterogenicity in the likelihood of different nucleotide substitutions, is used to explain the increasing or decreasing guanine-cytosine content (GC%) in DNA and is represented by microD, in agreement with Sueoka (1962, Proc Natl Acad Sci USA 48:582-592). The new method uses simulation to facilitate identification of significant A+T or G+C pressure as well as the comparison of directional mutation pressure among genes, even when they are translated by different genetic codes. We use the method to analyze the evolution of directional mutation pressure and nucleotide content of mitochondrial cytochrome b genes. Results from a survey of 110 taxa indicate that the cytochrome b genes of most taxa are subjected to significant directional mutation pressure and that the gene is subject to A+T pressure in most cases. Only in the anseriform bird Cairina moschata is the cytochrome b gene subject to significant G+C pressure. The GC% at nonsynonymous codon sites decreases proportionately with increasing A+T pressure, and with a slope less than one, indicating a presence of selective constraints. The cytochrome b genes of insects, nematodes, and eumycotes are subject to extreme A+T pressures (microD = 0.123, 0.224, and 0.130) and, in parallel, the GC% of the nonsynonymous codon sites has decreased from about 0.44 in organisms that are not subjected to A+T or G+C pressure to about 0.332, 0.323, and 0.367, respectively. The distribution of taxa according to the GC% at nonsynonymous codon sites and directional mutation pressure supports the notion that variation in these parameters is a phylogenetic component.

Biological Evolution

Molecular evidence for the inclusion of cetaceans within the order Artiodactyla.

The transition in the cetaceans from terrestrial life to a fully aquatic existence is one of the most enduring evolutionary mysteries. Resolving the phylogenetic relationships between Cetacea and the other orders of eutherian mammals may provide us with important clues to the origin of whales and may help us date the evolutionary transition to aquatic life. Previous paleontological and molecular evidence has indicated that cetaceans and artiodactyls constitute a natural clade within subclass Eutheria. Our present phylogenetic analyses of protein and mitochondrial DNA sequence data indicate that cetaceans are not only intimately related to the artiodactyls; they are in fact deeply nested within the artiodactyl phylogenetic tree; i.e., they are more closely related to the members of one suborder of artiodactyls, the Ruminantia, than either ruminants or cetaceans are to members of the other two artiodactyl suborders: Suiformes and Tylopoda. On the basis of the rate of evolution of mitochondrial DNA sequences and using paleontological reference dates for calibration, we estimate that the whale lineage has branched off a protoruminant lineage < 50 Mya. By implication, the cetacean transition to aquatic life is inferred to be a relatively recent evolutionary event.

Animals

Coral-host specificity of Red Sea Lithophaga bivalves: interspecific and intraspecific variation in 12S mitochondrial ribosomal RNA.

Comparison of 12S mitochondrial ribosomal DNA sequences was used to approach the question of species specificity between boring bivalves of the genus Lithophaga and their coral hosts. A 450-bp long fragment was amplified by PCR from 13 individuals belonging to five subgroups of Lithophaga bivalves. These subgroups are defined according to their coral hosts species, and they belong to three currently recognized species: L. lessepsiana (1 host), L. simplex (2 hosts), and L. purpurea (2 hosts). All bivalves were collected from corals growing within an approximately 200-m section of the reef of Eilat, Red Sea. Sequence variation between members of the same species inhabiting different hosts (30-32%) was found to be very similar to the variation exhibited between recognized species. These results, when interpreted together with previously published data concerning variations among Lithophaga subgroups, support the notion of a very high degree of species specificity between Lithophaga bivalves and their coral hosts in the Red Sea.

Animals

Towards a molecular resolution of the ordinal phylogeny of the eutherian mammals.

Reconstructing the evolutionary relationships among the orders of eutherian mammals entails the identification of a single true phylogenetic tree out of approximately 10(19) possible ones. The morphological and paleontological legacy to the field consists of numerous contradictory trees that are mostly devoid of binary resolution. With the introduction of molecular methodologies, several superordinal relationships have been identified, and in several instances a complete taxonomic revision was indicated. In this review, I present a summary of the phylogenetic affinities of the eutherian orders as revealed by molecular studies, and outline the differences between the molecular phylogenetic schemes and the phylogenetic trees produced through the use of morphological data. Questions of monophyly or paraphyly of the eutherian orders are also discussed. It is estimated that all but 10(9) of the 10(19) possible phylogenetic trees have been ruled out by molecular analysis, and that DNA and protein sequences with their potential to supply millions of phylogenetically useful characters will resolve the phylogeny of the orders of mammals into a consistently bifurcating tree in the not-so-distant future.

Animals

Structure and evolution of opossum, guinea pig, and porcupine cytochrome b genes.

We have sequenced the mitochondrial cytochrome b gene from the guinea pig, the African porcupine, and a South American opossum. A phylogenetic analysis, which includes 22 eutherian and four other vertebrate cytochrome b sequences, indicates that the guinea pig and the porcupine constitute a natural clade (Hystricomorpha) that is not a sister group to the clade of mice and rats (Myomorpha). Therefore, the hypothesis that the Rodentia is paraphyletic receives additional support. The artiodactyls, the perissodactyls, and the cetaceans form a group that is separated from the primates and the rodents. The 26 sequences are used to study the structure/function relationships in cytochrome b, whose function is electron transport. Most of the amino acid residues involved in the two reaction centers are well conserved in evolution. The four histidines that are believed to ligate the two hemes are invariant among the 26 sequences, but their nearby residues are not well conserved in evolution. The eight transmembrane domains represent some of the most divergent regions in the cytochrome b sequence. The rate of nonsynonymous substitution is considerably faster in the human and elephant lineages than in other eutherian lineages; the faster rate might be due to coevolution between cytochrome b and cytochrome c.

Amino Acid Sequence

The biochemical phylogeny of guinea-pigs and gundis, and the paraphyly of the order rodentia.

1. Molecular data indicate that caviomorphs (guinea-pig-like rodents) and myomorphs (rat-like rodents) are not monophyletic. 2. Rather, the evolutionary lineage leading to the guinea-pig may have branched off prior to the divergence among myomorphs, lagomorphs, primates, chiropterans, artiodactyls, and carnivores. 3. Thus, the guinea-pig lineage probably represents an ancient eutherian lineage, and should be conferred an independent ordinal status. 4. The gundis (Ctenodactylidae) also seem to have branched off before the divergence among myomorphs, primates, and artiodactyls, but after the divergence of the guinea-pig. 5. Therefore, the order Rodentia as defined at the present time is in all probability a paraphyletic group devoid of taxonomic validity. 6. Previous claims pertaining to large differences in the rate of molecular evolution between guinea-pigs and myomorphs may have been exaggerated in many cases as a result of the erroneous phylogenetic position attributed to the guinea-pig. 7. The average rate of amino acid replacement in the guinea-pig is comparable to that in the rat and the mouse. 8. Protein-coding genes of myomorphs and caviomorphs evole, on average, about two times faster than their counterparts in gundis and humans.

Amino Acid Sequence

Is the guinea-pig a rodent?

The guinea-pig (Cavia porcellus), traditionally classified as a New World hystricomorph rodent, often shows anomalous morphological and molecular features in comparison with other eutherian mammals. For example, its insulin differs from that of other mammals in anabolic and growth-promoting activities and in its capability to form hexamers. Indeed, the literature about the molecular evolution of guinea-pigs abounds in references to 'convergent evolution', 'extremely rapid rates of substitution', and 'unique evolutionary mechanisms'. These claims are based on the assumption that the guinea-pig is a rodent. Our phylogenetic analyses of amino-acid sequence data, however, imply that the guinea-pig diverged before the separation of the primates and the artiodactyls from the myomorph rodents (rats and mice). If true, then the myomorphs and the caviomorphs do not constitute a natural clade, and the Caviomorpha (or the Histricomorpha) should be elevated in taxonomical rank and regarded as a separate mammalian order distinct from the Rodentia. If, as suggested by recent data, the myomorphs branched off before the divergence among the carnivores, lagomorphs, artiodactyls and primates, then the new order would represent an early divergence in eutherian radiation.

Animals