(Below N is a link to NCBI taxonomic web page and E link to ESTHER at designed phylum.) > cellular organisms: NE > Eukaryota: NE > Opisthokonta: NE > Metazoa: NE > Eumetazoa: NE > Bilateria: NE > Protostomia: NE > Ecdysozoa: NE > Panarthropoda: NE > Arthropoda: NE > Mandibulata: NE > Pancrustacea: NE > Hexapoda: NE > Insecta: NE > Dicondylia: NE > Pterygota: NE > Neoptera: NE > Holometabola: NE > Diptera: NE > Brachycera: NE > Muscomorpha: NE > Eremoneura: NE > Cyclorrhapha: NE > Schizophora: NE > Acalyptratae: NE > Ephydroidea: NE > Drosophilidae: NE > Drosophilinae: NE > Drosophilini: NE > Drosophila [fruit fly, genus]: NE > Sophophora: NE > melanogaster group: NE > melanogaster subgroup: NE > Drosophila simulans: NE
LegendThis sequence has been compared to family alignement (MSA) red => minority aminoacid blue => majority aminoacid color intensity => conservation rate title => sequence position(MSA position)aminoacid rate Catalytic site Catalytic site in the MSA NYVGLIIVLSCLWLGSNASDPDDPLLVQLPQGKLRGRDNGSYYSYESIPY AEPPTGDLRFEAPEPYKQKWSDIFDATKTPVACLQWDQFTPGANKLVGEE DCLTVSIYKPKNSKRSTFPVVAHIHGGAFMFGAAWQNGHENVMREGKFIL VKISYRLGPLGFASTGDRDLPGNYGLKDQRLALKWIKQNIASFGGEPQNV LLIGHSAGGASVHLQMLREDFGQLAKAAFSFSGNALDPWVVQKGARGRAF ELGRNVGCESAEDSASLKKCLKSKSASELVTAVRKFLIFSYVPFAPFSPV LEPSDAPDAILTQDPREVIKSGKFGQVPWAVSYVTEDGGYNAALLLKERK SGIVIDDLNDRWLELAPYFLFYRDTKTKKDMDDYSRKIKEDYLGNQKFDI ESYSELQRLFTDILFKNSTQESLDLHRKYGKSPAYAYVYDNPAEKGIAQV LANRTDYDFGTVHGDDYFLIFENFVREVEMRPDEEIISRNFINMLADFAS SDNGVLKYGECAFKNNVGSEKFQLLAIYIDGCQNRQHVEFP
References
1 moreTitle: Latitudinal clines for nucleotide polymorphisms in the Esterase 6 gene of Drosophila melanogaster Coppin CW, Odgers WA, Oakeshott JG Ref: Genetica, 129:259, 2007 : PubMed
Previous studies have found non-neutral patterns of nucleotide polymorphism in the promoter and coding regions of Est6 in D. melanogaster. Coding region polymorphism peaks around two closely linked replacement differences associated with the EST6-F/EST6-S allozyme polymorphism. The promoter contains two common, highly diverged haplotype groups, P1 and P7, that differentially affect Est6 expression. Allozyme studies have also revealed latitudinal clines in EST6-F and EST6-S frequencies that recur across continents. Here we analyse nucleotide polymorphisms across the promoter and the region of peak coding sequence polymorphism in 10 Australian populations along a 25 degrees latitudinal gradient in order to examine the basis for the allozyme clines. As with the earlier studies, we find an excess of intermediate to high frequency variants in both the P1/P7 region and around the two EST6-F/EST6-S replacements in some populations. The two EST6-F/EST6-S replacement polymorphisms show latitudinal clines whereas the P1 and P7 groups of promoter haplotypes do not. However the strongest clines are for three co-segregating silent site polymorphisms in a 4 bp stretch at the 3' end of the sequenced region. Monte Carlo simulations show that the clines for those three sites can explain all others in the data but none of the others can explain those three. Thus the allozyme clines may not reflect selection on either the P1/P7 polymorphism or the two replacements previously associated with the EST6-F/EST-S difference.
        
Title: Nucleotide variation at the hypervariable esterase 6 isozyme locus of Drosophila simulans Karotam J, Boyce TM, Oakeshott JG Ref: Molecular Biology Evolution, 12:113, 1995 : PubMed
Esterase 6 (Est-6/EST6) is polymorphic in both Drosophila melanogaster and D. simulans for two common allozyme forms, as well as for several other less common variants. Parallel latitudinal clines in the frequencies of the common EST6-F and EST6-S allozymes in these species have previously been interpreted in terms of a shared amino acid polymorphism that distinguishes the two variants and is subject to selection. Here we compare the sequences of four D. simulans Est-6 isolates and show that overall estimates of nucleotide heterozygosity in both coding and 5' flanking regions are more than threefold higher than those obtained previously for this gene in D. melanogaster. Nevertheless, the ratio of replacement to exon silent-site polymorphism in D. simulans is less than the ratio of replacement to silent divergence between D. simulans and D. melanogaster, which could be the result of increased efficiency of selection against replacement polymorphisms in D. simulans or to divergent selection between the two species. We also find that the amino acid polymorphisms separating EST6-F and EST6-S in D. simulans are not the same as those that separate these allozymes in D. melanogaster, implying that the shared clines do not reflect shared molecular targets for selection. All comparisons within and between the two species reveal a remarkable paucity of variation in a stretch of nearly 400 bp immediately 5' of the gene, indicative of strong selective constraint to retain essential aspects of Est-6 promoter function.
        
Title: Conservation and change in structural and 5' flanking sequences of esterase 6 in sibling Drosophila species [published erratum appears in Genetica 1993;90(1):79-80] Karotam J, Delves AC, Oakeshott JG Ref: Genetica, 88 (1):11, 1993 : PubMed
Esterase 6 (Est-6/EST6) is the major beta-carboxylesterase in D. melanogaster and its siblings D. simulans and D. mauritiana. It is expressed in several tissues but its major site of expression is the sperm ejaculatory duct of the adult male. Although EST6 activity affects reproductive fitness, there are high levels of electrophoretic and activity polymorphism, at least within D. melanogaster and D. simulans. Here we present the nucleotide sequences of an Est-6 allele and its flanking regions from each of D. simulans and D. mauritiana and compare them with the published D. melanogaster sequences. As might be expected, replacement sites are significantly less divergent than exon silent sites in all comparisons, suggesting that selection is acting to maintain EST6 structure and function among the three species. Nevertheless, the ratio of the levels of replacement to silent site divergence is still much higher for Est-6 than for seven of ten other genes (including both isozyme-coding loci) for which comparable data have been published for these species. This is consistent with the high levels of EST6 electrophoretic polymorphism within D. melanogaster and D. simulans and implies that selective constraints against amino acid change are relatively weak for EST6. By contrast, comparisons involving promotor sequences show that the level of divergence in the first 350bp 5' of the gene is significantly lower than those for four of the six other loci for which comparable data have been published for these species. In particular, there are two perfectly conserved stretches (-1 to -158bp and -219 to -334bp) each over 100bp long included in this 350bp region. Thus the data suggest a relatively low level of selective constraint on the amino acid sequence of EST6 but a relatively high level of constraint on sequences affecting aspects of its expression.
Comparative analysis of multiple genomes in a phylogenetic framework dramatically improves the precision and sensitivity of evolutionary inference, producing more robust results than single-genome analyses can provide. The genomes of 12 Drosophila species, ten of which are presented here for the first time (sechellia, simulans, yakuba, erecta, ananassae, persimilis, willistoni, mojavensis, virilis and grimshawi), illustrate how rates and patterns of sequence divergence across taxa can illuminate evolutionary processes on a genomic scale. These genome sequences augment the formidable genetic tools that have made Drosophila melanogaster a pre-eminent model for animal genetics, and will further catalyse fundamental research on mechanisms of development, cell biology, genetics, disease, neurobiology, behaviour, physiology and evolution. Despite remarkable similarities among these Drosophila species, we identified many putatively non-neutral changes in protein-coding genes, non-coding RNA genes, and cis-regulatory regions. These may prove to underlie differences in the ecology and behaviour of these diverse species.
        
Title: Latitudinal clines for nucleotide polymorphisms in the Esterase 6 gene of Drosophila melanogaster Coppin CW, Odgers WA, Oakeshott JG Ref: Genetica, 129:259, 2007 : PubMed
Previous studies have found non-neutral patterns of nucleotide polymorphism in the promoter and coding regions of Est6 in D. melanogaster. Coding region polymorphism peaks around two closely linked replacement differences associated with the EST6-F/EST6-S allozyme polymorphism. The promoter contains two common, highly diverged haplotype groups, P1 and P7, that differentially affect Est6 expression. Allozyme studies have also revealed latitudinal clines in EST6-F and EST6-S frequencies that recur across continents. Here we analyse nucleotide polymorphisms across the promoter and the region of peak coding sequence polymorphism in 10 Australian populations along a 25 degrees latitudinal gradient in order to examine the basis for the allozyme clines. As with the earlier studies, we find an excess of intermediate to high frequency variants in both the P1/P7 region and around the two EST6-F/EST6-S replacements in some populations. The two EST6-F/EST6-S replacement polymorphisms show latitudinal clines whereas the P1 and P7 groups of promoter haplotypes do not. However the strongest clines are for three co-segregating silent site polymorphisms in a 4 bp stretch at the 3' end of the sequenced region. Monte Carlo simulations show that the clines for those three sites can explain all others in the data but none of the others can explain those three. Thus the allozyme clines may not reflect selection on either the P1/P7 polymorphism or the two replacements previously associated with the EST6-F/EST-S difference.
        
Title: Nucleotide variation at the hypervariable esterase 6 isozyme locus of Drosophila simulans Karotam J, Boyce TM, Oakeshott JG Ref: Molecular Biology Evolution, 12:113, 1995 : PubMed
Esterase 6 (Est-6/EST6) is polymorphic in both Drosophila melanogaster and D. simulans for two common allozyme forms, as well as for several other less common variants. Parallel latitudinal clines in the frequencies of the common EST6-F and EST6-S allozymes in these species have previously been interpreted in terms of a shared amino acid polymorphism that distinguishes the two variants and is subject to selection. Here we compare the sequences of four D. simulans Est-6 isolates and show that overall estimates of nucleotide heterozygosity in both coding and 5' flanking regions are more than threefold higher than those obtained previously for this gene in D. melanogaster. Nevertheless, the ratio of replacement to exon silent-site polymorphism in D. simulans is less than the ratio of replacement to silent divergence between D. simulans and D. melanogaster, which could be the result of increased efficiency of selection against replacement polymorphisms in D. simulans or to divergent selection between the two species. We also find that the amino acid polymorphisms separating EST6-F and EST6-S in D. simulans are not the same as those that separate these allozymes in D. melanogaster, implying that the shared clines do not reflect shared molecular targets for selection. All comparisons within and between the two species reveal a remarkable paucity of variation in a stretch of nearly 400 bp immediately 5' of the gene, indicative of strong selective constraint to retain essential aspects of Est-6 promoter function.
        
Title: Conservation and change in structural and 5' flanking sequences of esterase 6 in sibling Drosophila species [published erratum appears in Genetica 1993;90(1):79-80] Karotam J, Delves AC, Oakeshott JG Ref: Genetica, 88 (1):11, 1993 : PubMed
Esterase 6 (Est-6/EST6) is the major beta-carboxylesterase in D. melanogaster and its siblings D. simulans and D. mauritiana. It is expressed in several tissues but its major site of expression is the sperm ejaculatory duct of the adult male. Although EST6 activity affects reproductive fitness, there are high levels of electrophoretic and activity polymorphism, at least within D. melanogaster and D. simulans. Here we present the nucleotide sequences of an Est-6 allele and its flanking regions from each of D. simulans and D. mauritiana and compare them with the published D. melanogaster sequences. As might be expected, replacement sites are significantly less divergent than exon silent sites in all comparisons, suggesting that selection is acting to maintain EST6 structure and function among the three species. Nevertheless, the ratio of the levels of replacement to silent site divergence is still much higher for Est-6 than for seven of ten other genes (including both isozyme-coding loci) for which comparable data have been published for these species. This is consistent with the high levels of EST6 electrophoretic polymorphism within D. melanogaster and D. simulans and implies that selective constraints against amino acid change are relatively weak for EST6. By contrast, comparisons involving promotor sequences show that the level of divergence in the first 350bp 5' of the gene is significantly lower than those for four of the six other loci for which comparable data have been published for these species. In particular, there are two perfectly conserved stretches (-1 to -158bp and -219 to -334bp) each over 100bp long included in this 350bp region. Thus the data suggest a relatively low level of selective constraint on the amino acid sequence of EST6 but a relatively high level of constraint on sequences affecting aspects of its expression.