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A genomic survey of the fish parasite Spironucleus salmonicida indicates genomic plasticity among diplomonads and significant lateral gene transfer in eukaryote genome evolution Jan O. Andersson, Åsa M. Sjögren, David S. Horner, Colleen A. Murphy, Patricia L. Dyal, Staffan G. Svärd, John M. Logsdon, Jr., Mark A. Ragan, Robert P. Hirt, and Andrew J. Roger Additional file 3 Candidate LGT genes in the Spironucleus salmonicida genome survey. Clone Protein Classification SpESTZap1389 drug:H+ antiporter-2, DHA2 family Environmental information processing; membrane transport; major facilitator superfamily (MFS) Genetic information processing; folding, sorting and degradation; protein folding and associated processing Genetic information processing; folding, sorting and degradation; protein folding and associated processing Genetic information processing; replication and repair; other replication, recombination and repair factors Genetic information processing; unclassified SpESTC18 1 gZar958bT7 membrane protease subunit HflK 1 membrane protease subunit HflK SpESTC43 DNA glycosylase SpESTZap1340 tRNA (5-methylaminomethyl-2thiouridylate)-methyltransferase alanyl-tRNA synthetase aminoacylhistidine dipeptidase aminoacylhistidine dipeptidase aminoacylhistidine dipeptidase 5'-methylthioadenosine nucleosidase arginine deiminase ornithine carbamoyltransferase prolyl-tRNA synthetase sarcosine oxidase 27982008 1 SpESTC149 1 SpESTDH287 1 SpESTZap149 SpESTC31 SpESTC83 SpESTC221 27981826 1 gZar602gT3 gZap30gT3 1 sarcosine oxidase 27982357 threonine dehydratase SpESTZap946 1 gZap84gT3 1 SpESTZap588 27983189 27983404 SpESTDH325 histidinol-phosphatase (PHP family) alcohol dehydrogenase alcohol dehydrogenase alcohol dehydrogenase alcohol dehydrogenase alcohol dehydrogenase Tree# Metabolism; amino acid metabolism; alanine and aspartate metabolism Metabolism; amino acid metabolism; alanine and aspartate metabolism Metabolism; amino acid metabolism; alanine and aspartate metabolism Metabolism; amino acid metabolism; alanine and aspartate metabolism Metabolism; amino acid metabolism; arginine and proline metabolism Metabolism; amino acid metabolism; arginine and proline metabolism Metabolism; amino acid metabolism; arginine and proline metabolism Metabolism; amino acid metabolism; arginine and proline metabolism Metabolism; amino acid metabolism; glycine, serine and threonine metabolism Metabolism; amino acid metabolism; glycine, serine and threonine metabolism Metabolism; amino acid metabolism; glycine, serine and threonine metabolism Metabolism; amino acid metabolism; histidine metabolism Metabolism; carbohydrate metabolism; glycolysis / gluconeogenesis Metabolism; carbohydrate metabolism; glycolysis / gluconeogenesis Metabolism; carbohydrate metabolism; glycolysis / gluconeogenesis Metabolism; carbohydrate metabolism; glycolysis / gluconeogenesis Metabolism; carbohydrate metabolism; glycolysis / gluconeogenesis 1 (3) 3 #aa positions Monophyletic diplomonads 1 170 + 2 160 - DD 3 125 - DD 4 165 + DD 5 107 - 6 7 7 7 8 9 10 11 12 451 247 247 247 168 356 303 175 218 + + + + + + + + - 12 218 - 13 269 + 14 15 16 16 16 17 113 121 147 147 147 223 + + + + + - 4 4 Monophyletic diplomonads and parabasalids 5 Eukaryotetoeukaryote 6 LGT event 5 + + + + EH + + + DD EH EH + EH EH 23266714 SpESTZap404 18030018 gTol047bMF SpESTC275 gTor983gT3 fructose-bisphosphate aldolase, class II glucokinase glucose-6-phosphate isomerase phosphoglycerate mutase pyruvate kinase phosphoglycolate phosphatase SpESTC23 SpESTZap360 gZap797bT7 1 SpESTZap1028 1 gZap472bT7 SpESTC94 ribose 5-phosphate isomerase B transketolase acetyl-CoA synthetase (ADP-forming) hydroxyacylglutathione hydrolase hydroxyacylglutathione hydrolase putative pyruvate-flavodoxin oxidoreductase glycogen(starch) synthase carbamate kinase cystathionine beta-lyase cystathionine beta-lyase cytochrome c biogenesis protein CcmG, thiol:disulfide interchangeprotein DsbE glutamate synthase (NADPH) small chain hydroxylamine reductase tryptophanase tryptophanase NADH dehydrogenase NADH dehydrogenase NADH dehydrogenase NADH dehydrogenase NADH dehydrogenase cysteine synthase cysteine synthase fatty acid desaturase NAD(P)H dehydrogenase (quinone) long-chain fatty-acid-CoA ligase glycerol-3-phosphate dehydrogenase phosphatidylserine synthase tripeptide aminopeptidase adenine phosphoribosyltransferase deoxyguanosine kinase deoxyguanosine kinase myosin ATPase CTP synthase A-type flavoprotein A-type flavoprotein carotenoid isomerase carotenoid isomerase rubrerythrin rubrerythrin conserved hypothetical protein conserved hypothetical protein conserved hypothetical protein gZap510gT3 SpESTC239 2 gZap126bT7 2 gZap126gT3 1 SpESTC259 20530910 27983817 2 gTol313bMF 2 SpESTC204 gZal229bMF SpESTC219 SpESTDH113 1 11127702 1 gTor1143gT3 1 SpESTC115 1 SpESTZap1933 gZap471gT3 SpESTZap110 27983595 SpESTZap2148 gTor927gT3 gZar919bT7 SpESTC154 SpESTC254 SpESTC251 gTor125bT7 27982526 1 27984028 1 SpESTZap2025 1 SpESTC196 1 gZap998gT3 1 SpESTC282 1 SpESTC267 gTor1213bT7 gTor541gT3 gTor565bT7 Metabolism; carbohydrate metabolism; glycolysis / gluconeogenesis Metabolism; carbohydrate metabolism; glycolysis / gluconeogenesis Metabolism; carbohydrate metabolism; glycolysis / gluconeogenesis Metabolism; carbohydrate metabolism; glycolysis / gluconeogenesis Metabolism; carbohydrate metabolism; glycolysis / gluconeogenesis Metabolism; carbohydrate metabolism; glyoxylate and dicarboxylate metabolism Metabolism; carbohydrate metabolism; pentose phosphate pathway Metabolism; carbohydrate metabolism; pentose phosphate pathway Metabolism; carbohydrate metabolism; pyruvate metabolism Metabolism; carbohydrate metabolism; pyruvate metabolism Metabolism; carbohydrate metabolism; pyruvate metabolism Metabolism; carbohydrate metabolism; pyruvate metabolism 18 19 20 21 22 23 300 192 412 184 205 116 + + + + 4 + - 24 25 26 27 27 28 136 196 121 164 164 167 + + + + + + Metabolism; carbohydrate metabolism; starch and sucrose metabolism Metabolism; energy metabolism; nitrogen metabolism Metabolism; energy metabolism; nitrogen metabolism Metabolism; energy metabolism; nitrogen metabolism Metabolism; energy metabolism; nitrogen metabolism 29 30 31 32 33 111 273 157 182 110 + + + Metabolism; energy metabolism; nitrogen metabolism 34 243 + Metabolism; energy metabolism; nitrogen metabolism Metabolism; energy metabolism; nitrogen metabolism Metabolism; energy metabolism; nitrogen metabolism Metabolism; energy metabolism; oxidative phosphorylation Metabolism; energy metabolism; oxidative phosphorylation Metabolism; energy metabolism; oxidative phosphorylation Metabolism; energy metabolism; oxidative phosphorylation Metabolism; energy metabolism; oxidative phosphorylation Metabolism; energy metabolism; sulfur metabolism Metabolism; energy metabolism; sulfur metabolism Metabolism; lipid metabolism; androgen and estrogen metabolism Metabolism; lipid metabolism; biosynthesis of steroids Metabolism; lipid metabolism; fatty acid metabolism Metabolism; lipid metabolism; glycerophospholipid metabolism Metabolism; lipid metabolism; glycerophospholipid metabolism Metabolism; metabolism of other amino acids; glutathione metabolism Metabolism; nucleotide metabolism; purine metabolism Metabolism; nucleotide metabolism; purine metabolism Metabolism; nucleotide metabolism; purine metabolism Metabolism; nucleotide metabolism; purine metabolism Metabolism; nucleotide metabolism; pyrimidine metabolism Metabolism; unclassified Metabolism; unclassified Metabolism; unclassified Metabolism; unclassified Metabolism; unclassified Metabolism; unclassified 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 52 53 54 55 56 57 57 58 58 59 60 61 471 160 164 105 315 155 303 155 200 214 143 127 682 185 168 214 154 182 182 135 511 383 168 211 211 173 173 144 199 160 + 4 + 4 + + + + + + + + + + + + + + + 2 (3) + + + MB EH EH EH EH DD DD + + GZ gTor916gT3 gTor930bT7 gZap260bT7 gZap950bT7 gZar587bT72 gZar847bT7 gZar96bT7 1 gZar978gT3 1 gZar789bT7 Sp11orf4 SpESTZap1004 SpESTZap1248 1 SpESTZap1299 1 SpESTZap1913 conserved hypothetical protein conserved hypothetical protein conserved hypothetical protein conserved hypothetical protein conserved hypothetical protein conserved hypothetical protein conserved hypothetical protein conserved hypothetical protein conserved hypothetical protein conserved hypothetical protein conserved hypothetical protein conserved hypothetical protein conserved hypothetical protein conserved hypothetical protein 62 63 64 65 66 67 68 69 69 70 71 72 33 33 175 163 123 111 124 137 147 185 185 586 197 118 110 110 + + + + + + + + + + + 1 ) The phylogenetic analyses of these genes indicate that they likely have undergone duplications after the putative LGT events. ) These pairs of sequences with identical annotations correspond to different parts of the protein and could represent the same gene in the genome. 3 ) Phylogenetic trees for all candidate LGT genes are shown in Additional files 4-6. 4 ) G. lamblia homologs for these proteins are present in the similarity searches and most likely are the closest relatives to the S. salmonicida sequences. However, the G. lamblia homologs are absent from the phylogenetic trees listed since the automated generation of datasets [1] reached the maximum number of sequences to include (200) and only selected a subset of the available homologous sequences in the databases. 5 ) A previous phylogenetic study of alanyl tRNA synthetase has both shown diplomonads and parabasalids to be monophyletic, and gene transfers between eukaryotic lineages [2]. These cases are not observed in the presented tree since the Trichomonas, Entamoeba, and ciliate homologs were excluded due to the reasons outlined above. 6 ) The putative recipient and/or donor eukaryotic lineages are indicted by DD (Dictyostelium discoideum), EH (Entamoeba histolytica), GZ (Gibberella zeae), and MB (Mastigamoeba balamuthi). 2 References 1. 2. Frickey T, Lupas AN: PhyloGenie: automated phylome generation and analysis. Nucleic Acids Res 2004, 32:5231-5238. Andersson JO, Sarchfield SW, Roger AJ: Gene transfers from Nanoarchaeota to an ancestor of diplomonads and parabasalids. Mol Biol Evol 2005, 22:85-90. 3 (3)