Opsin evolution: Peropsin phyloSNPs

Introduction to PhyloSNPs in Peropsin Evolution

PhyloSNPs are defined as single coding residue changes at a site (1) strictly conserved ancestrally over great branch lengths but (2) switching in an ancestral stem to a new residue that in turn is (3) strictly conserved in all descendent extant species. A certain amount of excursion from perfect conservation can be admitted (reduced alphabet, disproportionately attributable to sequencing error) and even low levels of homoplasy (where the ancestral value reoccurs in a later diverging a species or two. Peropsin (or RRH) has just a handful of them at its 337 positions, the number depending on quality cutoff. Most of these occured in the earliest mammals.

PhyloSNPs are thus similar to synapomorphies but are adapted to protein molecular evolution and its fixed alphabet of 20 distinct, unambiguously determinable charcters. Because phyloSNPs are fixed both before and after the stem event, they greatly enrich for defining characters of the stem clade (and its complement) -- in effect they answer the question, what makes a mammal a mammal. Synapomorphies (eg vertebral column) don't have this requirement on the clade complement -- ie sea urchins don't have notochords.

PhyloSNPs differ from conventional SNPs familiar from human variation studies in their phylogenetic depth. Differences in the human population or even human/chimp speak to a much shorter time scale with few or no intermediate divergence nodes (to extant or sequencable extinct species). Consequently these variations are not readily interpretable nor adaptive, whereas high quality phyloSNPs in stem placentals (at the current level of 37 mammalian genomes and 13 non-mammals) have accrued over a billion years of branch length during which the phyloSNP residue has been conserved in all species (either as the clade-specific change or ancestral value). Consequently phyloSNPs must represent adaptive change at the residue in question. However phyloSNPs don't capture everything: features such as glycosylation sites and disulfides can evolve out of 'nothing' (quite variable sequence lacking a specific alternative ancestral function).

The timing of phyloSNP establishment must be considered in relation to timing of gene duplications within the ambient gene family. If the phyloSNP precedes the duplication, then it may well descend into descendent clades of the gene tree. These phyloSNPs might appear to be independent events if the individual paralogs are considered in isolation. Opsins are a moderately sized gene family with both paralogous and orthologous phyloSNPs (though so far peropsin only seems to have the latter).

Interpretation of phyloSNPs is not so easy from bioinformatics alone. In opsins, a great deal of structural and experimental information is availble. For example, a peropsin douple phyloSNP in adacent N.S columns at positions 211-213 might appear to be an ancestral glycosylation site lost in a coordinated way in the post- marsupial stem but before atlantogenata diverged, perhaps suggesting a shift in subcellular location in placental mammals. However, the site in fact is on the cytoplasmic side and consequently cannot have been a glycosylation site to begin with.

Note the species are listed below in more or less phylogenetic order (up to an arbitrary choice of human at the top). More precisely, various admissible symmetry operations can result in some swapping of order (eg of marsupials or murid rodents). These reorderings are limited to superordinal clades -- no matter what combinations of tree reorderings (leaving human as fixed point), mouse always stays above dog (ie, Euarchontoglires stay above Laurasiatheres.)

PhyloSNPs are required to be invariant (stay a connected vertical chain) under the action of this finite abelian subgroup of the species tree's permutation group. In other words, they are defined on the quotient tree under this action (rather than on the tree itself). For example, if a phyloSNP continues into the marsupial macEug but not into monDom, it would not be a phyloSNP because swapping order breaks the vertical continuity. Thus, to the extent the assumed tree is valid, there is no arbitrariness in the phylogenetic ordering of the y axis (orthology). PhyloSNP is a strong condition.

In a cladesheet that considers a single phyloSNP, the x-axis is the gene tree. That too has a distinct fixed topology and corresponding action of a separate abelian group of rearrangements. A paralogous phyloSNP must be well-defined here too. In opsins, the counterion at position 113 has both phyletic and paraphyletic phyloSNPs with respect to the gene tree, with the latter still being valid phyloSNPs with respect to the species tree.

It's worth noting the tremendous distillation of sequencing effort that goes into defining a single phyloSNP. First 50 large vertebrate genomes must be carefully sequenced. Of that data, only one-fiftieth codes for protein. Of that, only one-hundredth is a phyloSNP. Of those perhaps one-tenth are high enough quality and on a stem of interest. Of those perhaps one-tenth are stem events on a gene tree (which for opsins involves comparison of 15 proteins). Of those perhaps two-fifths are interpretable. Overall, a good phyletic phyloSNP represents over a million-fold reduction of raw data. That's precious signal relative to the background level of near-neutral noise and the near-meaningless drift in protein evolution.

PhyloSNPs in vertebrate peropsins

PhyloSNPs emerge from aligning 48 peropsins from cartilaginous fish to human: they are columns towards the bottom of ancestral residues that continued past various divergence nodes in a conserved manner, only to change at a more recent node in its descendent clade (here typically mammals), in contrast to the ancestral value which continues to the present day in other descendent species.

PhyloSNPs can be classified in various ways, amounting to horizonal sorts of the cladesheet (a spreadsheet with topological ordering of rows) by the criteria below. When phyloSNPs become available in all opsin paralogs (indeed all GPCR receptors), it may be instructive to extend sorting across the entire gene family to identify structural hotspots and coldspots of change.

• -- derived amino acid (synapomorphy)
• -- phylogenetic depth: fish, land, mammal, theria, placental
• -- overall assessed phyloSNP quality: A best, B some noise, C some homoplasy
• -- physiochemical severity of change
• -- position within sequence
• -- coding exon number within gene
• -- relative order by phylogenetic depth
• -- relative order within linear sequence
• -- topology: membrane, cytoplasmic, or extracellular
• -- secondary structure: alpha helix, beta sheet, coil

Peropsin exhibits 12 phyloSNPs of varying degrees of quality and interest. The first four cannot yet be fully evaluated because no outgroup data is available from cartilaginous fish or lobe-finned fish -- these phyloSNPs may only have relevence in defining key events within ray-finned fish. This will be resolved by elephantfish and coelocanth genome projects.

In the second group, the high-quality glycine to serine phyloSNP occured in a transmembrane segment sometime after platypus but before marsupial divergence. While this is a commonly observed change (high PAM coefficient in the Dayhoff matrix), those statistics are averaged over thousands of mostly cytoplasmic proteins and so have little bearing on specific positions in specific integral membrane proteins.

In the third group, the alanine to proline A --> P change at position 59 in a cytoplasmic loop could potential result in a kink so more rigid conformation with potential conformational and functional implications. This residue position has been discussed before as a signature histidine uniquely characteristic of SWS2 opsins. Two independent phyloSNPs at the same position at vastly different evolutionary times shows the amino acid at position 59 -- proline, alanine, or histidine -- is adaptively exploitable in different ways yet always under strong selection. Peropsin is not closely related to SWS2 in the opsin gene tree, meaning these events have not descended down from a gene duplication.

The asperagine to basic histidine change at position 211 membrane boundary is temporally coupled with a serine to larger but similar threonine change at neighboring cytoplasmic position 213, possibly as coevolutionary change. (In opsins, because the 3D structure is available, we can also consider coevolutionary changes of residues that are adjacent not in the linear sequence but only in tertiary structure.)

PhyloSNPs in peropsin are readily located in the overall alignment of 230 opsins by web browser search with a few flanking residues. The conservation of residue position in the overall scheme of opsin structural and functional evolution can contribute to the interpretation of individual peropsin phyloSNPs (as we have seen in the DRY to GRY phyloSNP of RGR opsin).

When this is done for the A --> P change at position 59, a very interesting result emerges: proline is actually the deep ancestral value in all other classes of opsins including all invertebrate opsins (with the exception of the SWS2 mentioned above). Consequently the sequence of events in peropsin must have been: P --> A --> P with the first change predating vertebrates and the 'reversion' postdating marsupial divergences. Indeed, the peropsin story becomes very muddled in amphioxus and sea urchin homologs because of gapping and weakened alignment. Peropsin is undoubtedly an ancient GPCR persisting from early eumetazoa yet it cannot be located outside of deuterostomes -- perhaps this change was critical in recruiting it to photoreceptor functionality.

.......................*......
PERa_braBe  P  KFRSLR-SPTTMLLV
PERa_braFl  P  KFRSLR-SPTTMLLV
PERb_braBe  A  KWRQLCRKAPNLLVI
PERb_braFl  A  KWRQLCRKAPNLLII
PER1_strPu  S  RYGTFRKRSVNILLM
PER2_strPu  S  RYRTFRKRSINLLLI

We've previously considered the case of the glutamate counterion at position 113 of cone/rod opsins but tyrosine ancestrally in most other opsins including peropsin. This locus thus exhibits a phyloSNP fixed prior to a series of gene family expansions in the stem of parapinopsin PPIN as well as later phyloSNPs localized within individual gene families. This suggests that considering paralogs helps even in evaluating orthologs to the extent that only a limited class of residue positions have phyloSNP potential, with that potential being used repeatedly at different junctures.

The phyloSNP at position 59 (human peropsin numbering) is quite interesting when considered across the entire gene family. The cladesheet is not fully populated -- only sequences from the opsin reference collection and the UCSC genome browser 28-way track are shown. (That data could be doubled by using genomes-in-progress stored at the NCBI trace archives.)

What is notable about this residue is not only the P --> A --> P phyloSNP in peropsin but also the P --> H --> gene loss seen in SWS2. This histidine is a departure from the expected proline that must have been present in the ancestral cone opsin protein that experienced considerable expansion between amphioxus and lamprey divergences. Details of that era are largely invisible to us for want of extant species or plausible sources for extinct species dna. Melanopsin 2 is also of interest for its leucine in early diverging species; this gene must be carefully separated from possibly similar extra melanopsins resulting from the fish-specific whole genome duplication.

The opsin family is somewhat restricted in terms of illustrating gene family aspects of phyloSNPs because of fairly massive gene loss (7 of 15) in mammals. We see here however that peropsin, neuropsin, and RGR (which are sometimes lumped as 'outsiders' to mainstream visual opsin development) have no deep connection at position 59. A more systematic evaluation of this requires full scale curation of the full set of opsin genes in vertebrates.

Alignment of 48 peropsins

..........................................................................................................1.........1.........1.........1.........1.........1.........1.........1.........1...1
................1.........2.........3.........4.........5.........6.........7.........8.........9.........0.........1.........2.........3.........4.........5.........6.........7.........8...8
.......1234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234
.......eeeeeeeeeeeeeeeeeeeeeeeeeeMMMMMMMMMMMMMMMMMMMMMMMccccccccccccMMMMMMMMMMMMMMMMMMMMMMMMMMeeeeeeeeeeeeeeMMMMMMMMMMMMMMMMMMMMcccccccccccccccccccMMMMMMMMMMMMMMMMMMMMMMMMeeeeeeeeeeeeeeeeeeee
exon...1111111111111111111111111111111111122222222222222222222222222222222222222222222222222222222222222233333333333333333333333333333333334444444444444444444444444444444444444444444444444444
marker........glyc....................................................................................diS.......................DRY................................................diS.........
.......................y..............................xx.........x...............xx..........x...............y.........................y.......................................................
opsins M........S..................A.Y...........N...L...v..K.LRTP.N.I..NLA..D......Gy.......LhG...FG..GC..........G......L.V.A.DRY..VC.P....R.........I...W......A..P..GW..Y.PD.....C.I.......
Consen M.............e..S.Fsq.EHnIVA.YLI.AG.iSi.SN..VLgiF.k.keLRTpTNa!IiNLA.TDIGVs.IGYPMSAASDl.GsWkFG..GCQIYAgLNIFFGMaSIGLLTvVA.DRYLTiC.Pd.Gr.mt...Y..mIl.AW.N..FWa.mP..GWA.YAPDPTGATCTiNWR.ND.
homSap MLRNNLGNSSDSKNEDGSVFSQTEHNIVATYLIMAGMISIISNIIVLGIFIKYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQVYAGLNIFFGMASIGLLTVVAVDRYLTICLPDVGRRMTTNTYIGLILGAWINGLFWALMPIIGWASYAPDPTGATCTINWRKNDR
panTro ........................................................................................................................................................................................
gorGor ........G................................................................................................................................................L..............................
ponPyg ....................................I..............................................................................................I....................................................
nomLeu ........................................................................................................................................................................................
macMul ........................................L..........................................................................................I...........M........................................
papHam ........................................L..........................................................................................I...........M........................................
calJac .............................S..........L...............................................................................A..........I......S...IM................V.......................
tarSyr ..K.D...................................L.........V................S...............................I............................R..I.........V.M............S...A...T..................A
otoGar ........N...............................L..............................................H...........I............................R..I.......S...M.....V..........T......................A
micMur                                                  I....................M.......R..I......H..V.M.....V......V...T...G..................A
tupBel ..GSDA....NLG...S.A..........A..LT..V...L...V.....VTH...............F..................H.R....H....I..........................L.R.A.....GSS..AAM.....L......A...A...G..................A
musMus ..SEASD...G.RS.G-....R...SVI.A...V..IT..L..VV.................V.....F..................H......H....I..........V.........M......SC............LSM....................................N..T
ratNor ...DA.D...G.GS.G-...TKS..S.I.A...V..I...L.....................V.....F..................H......H...............V.........L......SC........G...LSMV...............V......................T
speTri                                                                                            
dipOrd .....V....G.R....................T..V...L..L........................L....................R.........I....................I.......H..I..G...R..VTM.......................................T
cavPor ...HS........................A...L..L...L.......................M...L..................H........................................R..I.....SHS.V.M.......................................V
oryCun ......S....F.H...................L......L..L........................F..................H...........I....................M.......H.........R..L.......V..........A......................T
ochPri ...H......EA.V.A.............A...L......L..L........................F..................H...........I....................M.......Q..I......H..F.M................V......................K
canFam ......D..T...................A...T..I...F..L.......................................................I....................M.......S..T..........SM.....L.................................A
felCat ......D......................A...T.                                I....................M.......S.NS..........SM.....L.................................A
bosTau ...........C...N.............A...T..V...L...........F..................................H...........I............................H..A.....A....SM.....T.................................V
turTru ..................T..........A...T.....VL..............................................H...........I............................C.GA..........SM.....F..........V......................V
susScr ..LMD...N................S...A...T......L...V.......H...............A..................H...........I............................R.EA..........SM.......................................V
vicVic ...............Y......A......A...T......L...V.......H..............................................I....................M.......R..A..........SM............SV......G...............Q..V
equCab .............................A..........L.........V.......S..............................R.........I....................L.....T.R..A......S..TSM.....V..........V...N..................A
myoLuc ..K...S....F............................L..V.......T...................................H...........                 ..K...H...SM.......................................V
pteVam ..........N......................I......L.........F................................................                 ..........IM.......................................A
sorAra ..E.RSD...GPTP.G.A.L.R...RL..A...V..V...L..TA.............R.....M...L..................H...R..H......................L........L.R..A..S....S.V..........F.......L.........M.........G..V
eriEur ..E.....G...E................A...V.                                I............................R.HT..S.SA.S..AM.....L..........S...G...............E..G
loxAfr ....S.D.........A.......................L..............................................H.R.....T...I............................H.HI.....S...VSM.........L...L..T......................A
proCap ...............E.............A.....                                I..........S.........I.......H.N.                          
echTel ..K..E.T..G.HS.E......A..R...S...T......L..F..................M........................H.............................A..........H..R.....S...V.M........FL...L.V....................E..A
dasNov .....T.......D......................I...F......S.............T.....................................I..........T.........M.......R..T.....I....SM................A.....................NI
choHof                  .I...L...V................TV....................................I............................H........I....SM.............L.VT......................G
monDom .FK..SVKTLAPEK.GP....PI..K...A...T..V...V..V......V...A...A..T.................................D...I.................A..I.......Q..L.G...SYN.TLM..T..V..F.......V...G..................V
macEug .FQ.DS---LEPEK.SY....P.......A...T..V...P.........V.......A..T.....................................I.................A..I.......Q..L                          
ornAna .R..DSA.LLE.EHH.R.A....D.....A...T..IM..V..V......V.FE....A................G...........H......H....I..........S.................R.AI..K..RSN.TAM..A..M..F...S..LL......S...............A
galGal .HW.DSA...E.DA.AH...T........A...T..V...F...V.....V.......A.........F......G...........H.......T...I..A.........................R..I......RN.AA...A....AV...S..TV...G..S........A......T
taeGut .HW.DSS...E.DD.AH.A.T........A...T..V...F...V.....V.......A.........F......G...........H.......T...I..A.........................R..I......RS.AT...A....AV..SS..TA...............V......V
anoCar .FL.DSA...E.DD.PH.A...A......A...T..V..LL...V.....V.......A.........F......G...........H.......T...I..A.................I.......K.HI.S.L.ATN.TT...A....A....S..VV...............V......T
xenTro .AGTGTV.I..ASS.VH.....S......A...T..V...L.........V.......A.........F......G...........H.......V...I....................I.......R..I...ISGRH.TAM..A....AV..SV..VV..S...................A
danRer .ESGL.NV.AETVYGEK.A.T........A...T..V..LS...V..LM.V.FR....A.........F.....AG...........H.......M...I..A.................I.......R..I.QKL..RS.TL..VA..L.AV..SS...V...G...............N..V
takRub ..KVFSLVNISNEV.GK...T.W......G...I..V..LT...V..LM.V.F.....A..F......F.....AG..........IH......HT...I..A.................I...I...R..I..K..VQS.NL...A..L.AV..SS..VV...A...............Q.NV
tetNig ..KVFSLVN.SNDV.GK.A.T.W......G...T..I..LT...V..LM.V.F.....A..F......F.....AG..........IH......HT...I..A.................I.......R..I..K..VQS.NL..AA..L.AV..SS..VV...A...............Q.NA
gasAcu .DPEVNVTDDVTLYGGK.A.T.L......G...T..V..LF...V..LM.W.F.....A..F......F.....AG..........IH...........I..A.................I.......R..I.QK..MQS.NL...A..L.AV..SS..VV...................Q..A
oryLat .DSAVNVSDAAAPYGGK.A.T.L......G...T..V..LS..VL..LM.V.FR....A..I......F..V..AG..........IH.......T...I..A.................I.......R..L.QK..MQS.NL...A..L.AV..SS...V...G...............Q..V
calMil                                                                                            
homSap MLRNNLGNSSDSKNEDGSVFSQTEHNIVATYLIMAGMISIISNIIVLGIFIKYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQVYAGLNIFFGMASIGLLTVVAVDRYLTICLPDVGRRMTTNTYIGLILGAWINGLFWALMPIIGWASYAPDPTGATCTINWRKNDR
Consen M.............e..S.Fsq.EHnIVA.YLI.AG.iSi.SN..VLgiF.k.keLRT.TNa!IiNLA.TDIGVs.IGYPMSAASDl.GsWkFG..GCQIYAgLNIFFGMaSIGLLTvVA.DRYLTiC.Pd.Gr.mt...Y..mIl.AW.N..FWa.mP..GWA.YAPDPTGATCTiNWR.ND.
opsins M........S..................A.Y...........N...L...v..K.LRT..N.I..NLA..D......Gy.......LHG...FG..GC..........G......L.V.A.DRY..VC.P....R.........I...W......A..P..GW..Y.PD.....C.I.......
.......................y..............................xx.........x...............xx..........x...............y.........................y.......................................................

.......1....1.........2.........2.........2.........2.........2.........2.........2.........2.........2.........2.........3........3..........3.........3.......3
.......8....9.........0.........1.........2.........3.........4.........5.........6.........7.........8.........9.........0........1..........2.........3.......3
.......5678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678
.......eeeeMMMMMMMMMMMMMMMMMMMMMMMMccccccccccccccccccccccccccccMMMMMMMMMMMMMMMMMMMMMMMMeeeeeeeeMMMMMMMMMMMMMMMMMMMMMMMMMcccccccccccccccccccccccccccccccccccccccc*
exon...5555555555555555555555555555555555555555555555555555555566666666666666666666666666666666666666666666666666666666666677777777777777777777777777777777777777
..........................................................................................................K......................................................
.................................x.y....................................................................................................y...x....................
opsins ...Sf........Fi.PL.V.F.CY.......K....................VT.M.iiMI..FL..W.PY..V.............P....I...FAK.S..YNP.IYV..NK.FR........C...........................
Consen SFVSYTM.Vi.iNF..PL.VMfYCYY.V.........s.c....n.DWs.Q.DVTKMSvIMI.MFL.AWSPYSIVCLWasFGDPk.Ipp.mAIiAPLFAKSSTFYNPCIYV.ANKkFRrA..aM..Cqt.q.......lPM......l......
homSap SFVSYTMTVIAINFIVPLTVMFYCYYHVTLSIKHHTTSDCTESLNRDWSDQIDVTKMSVIMICMFLVAWSPYSIVCLWASFGDPKKIPPPMAIIAPLFAKSSTFYNPCIYVVANKKFRRAMLAMFKCQTHQTMPVTSILPMDVSQNPLASGRI*
panTro .........................................................................................................................................................*
gorGor ......V.................................................                              .....................................*
ponPyg .........................................................................................................................................................*
nomLeu .................................Y....................................................................................K.......WPN.....G.............T..K.*
macMul ...................................A..........E..........................................................................................................*
papHam ...................................A..........E...............................................................M..........................................*
calJac .......A......................F......................................................N...A..................................L............V....I..........*
tarSyr T......A...................A.Q.....VA.N.V..........V....                              .....F..L....Y.A..A..S...N......T..KN*
otoGar                             .......................L.........S...............................F.........A.A........I.......R..*
micMur ...........V..V...M..........Q...R..ATN............V....................A.........N.E....S.....................I.........F.........A......F..G......P...T*
tupBel .......A...V..V......S...VL.ARA.AG.AA.S.S.HRC....E.V.......L..L.....................QR...A...V.................L......K..C.........A.S...V...AS.PR...PA.V*
musMus .......M..VV................SR.LRLYAA....AH.H...A..A..........L...L............C..N......S.....................A.H....K.........P.LAV.EP.T....MP.SS..PV..*
ratNor .......M..VV................SQ.MRLSAA.N..TH.....AH.A.......M..L...L......V.....C..N......SL....................A......K..F..L...P..A..EP.T.A.G.PHS...PA..*
speTri ...........V.................R.....AA.N..AY........V..........L...................N......S.....................A...R.....F.........A.....V.......S.R.....*
dipOrd                             ...V..L......................E......................................L......A.....................*
cavPor A.................A......L.I.RA.RR.VAG.RPPN.SG.....V.......V..L.....................RR.S.S.....................I.........F...Q.....AV..A......A..S.......*
oryCun ....F..A......V..............Q...Q.RA.....Y........V..........F..........................A.....................A...R.....F.........A.....V..........P..I.*
ochPri ....F..A..MV..V...........Y..Q......A....K.........V..........L.....................Q....S.....................A...RS....F......IP.AK....LS.R....S..S...T*
canFam F...F......V.................R...C....N...Y........V..........F..........................S............................K.IF.........A..G.................N*
felCat ....F.........N..............QT..C.D..N..GY......N.V....                              ..K..F.....ENR.P................T...K*
bosTau .......M.V...................Q.....G.NN...Y........V..........L..........................S.....................I.................T.A.....V.....P....T..KV*
turTru .......M.V........IM.........R.....A..N...Y........V..........L..........................SV..L.................I...................A..ME.......P....T..KV*
susScr .........V.V......I..........Q...S.A..N..AY........V..........L................................................I...................A..LE.T.....P........V*
vicVic .......M.V........I..........R...CRA..N...Y........V..........L...L......................S.....................I...................A..M........P....T...L*
equCab .......A.V...................R.M.R.P..N..QY..T.....V..........F..........................S...............................F........RA...........P..Q......*
myoLuc ....F........................R...CRA..N...Y.....A..V..........F..........................S...............................F...........TTM.F.....P....T....*
pteVam ....F.............A..........R...C....N...Y........V..........F..........................S...............................F......D..S.....V.....P....T....*
sorAra .........V.V..VM.....A......IR.L.C...HSSPGH.DG...S.V.......V..LA..A.........F............S..V............................S..LT.RAQGA..AA.T....AAHS.Q....N*
eriEur ..............M...A..L.......R.M.C..ARS.A.YVHG.....V..........L..........V...............S...M.................L.........F.........A....NT....IP.K-.D.R.N*
loxAfr ..........V...V...A..........R...R..A.N.A.Y........L..........L....................S.....S...............................F.........AE...C....N.......A...*
proCap ......V...MV..V...A........I.....C.AARN.A.P.C......L..........L...M.................T....S.........................................AV...N....T....SS.....*
echTel .........VG...V...A..........QT..RQ.A.NGA.N.H......L....                              .....F.LLQ..PQEARR.......N.....M....L*
dasNov .........V....L...A..L.......R......K.N.S.YF.....N.V..........S.....................E....S..............................IF..L......A...M.....N..E........*
choHof .........VV...V..VA........I.R...RRNS.N.S.Y........V..........L...L......................S............................TI.F..L......AV........N..E........*
monDom ..........T...AM..G.......N.SQKM.QYSP.N.PDHI.....N.VA......V..L...L..................E...A...V.................A........IS..IR.....S..ISNA...N*     
macEug ..............VM..V.......N.S.KM.QY.R.S.P.HI.....N.V..........L...L......V...........E...A.....................A........IS..MR.E...S...SNA..LNLT*    
ornAna ..I........V..A...I.......N.SKAMRQYPA.RVL.N..I...E.V.......V..L...M...........S........S.AV..M............................S.VQ....REITI.DV...NR.RS..TL  
galGal .......S...V..V...........N.SRTM.QY.S.N.L..I.M.....V.......V..V...............S........S.A.....................I........I...VR...R.EITISNA...T..LSA.T.  
taeGut ..I....S...V..V...........N.SRTM.QYAS.N.L..I.I.....V.......V..I...............S........S.A.....................I........I...VR...R.EITINNA...S...SA.T.QNS*
anoCar .......S...V..VI..S.......N.SKTM.YYMRNS.L.NI.I.....V.......V..I...L...........S........S.A...V.................I...R....I...IR...R.EITINNV...S...STI.  
xenTro .......S.V.V..V...M.......N.SRTM.GYGSRSSLGGI.A.....T......MV..V...............S.....R....A.....................I........I.S.VQ.KSR.EVTLDNHF..N...ST.TT* 
danRer ..........TV...I..S.......N.SATV.RFKA.N.LD.I.M.....M..........V...A.................Q...A........L.............I........IIG.IR...R.RVTINNQ...MA.SV..NP* 
takRub ..I....A..GV..ML..F.......N.SVTVNRCK..N.LDDI.IE..E.M......LV..I......................A..A......................I........IIG.IR...R.Q.TINTEI..TT..QTATQ* 
tetNig ..I....A..SV...L..F.......N.SVTV.QYKANN.LDNI.IE..E.M......IV..I......................T.SA......................IT.....Q.IIG.IR...R.QITINTDI..TA..QT.TQ* 
gasAcu ..I....A...V..VL..SA......N.SATV.RYKA.N.LD.A.I.....M......IV..I......................T..A......................I........IIG.VR...R.RITIN.QV..TT..Q..TQ* 
oryLat .......A...V..V...........N.SATV.RYKA.N.LD.I.I.....M......IV..V..........M...........T..A......................I........IIG.IR...R.RITISTQV..TI..Q..TQ* 
calMil L..........V..V...S...F...N.SKTMSRFIS.PSP.NI.L.....L.......V..V...L...............N..L...A.....................I......K.IM..IC..NR.EITINHT...TI.RV..TE* 
homSap SFVSYTMTVIAINFIVPLTVMFYCYYHVTLSIKHHTTSDCTESLNRDWSDQIDVTKMSVIMICMFLVAWSPYSIVCLWASFGDPKKIPPPMAIIAPLFAKSSTFYNPCIYVVANKKFRRAMLAMFKCQTHQTMPVTSILPMDVSQNPLASGRI*
Consen SFVSYTM.Vi.iNF..PL.VMfYCYY.V.........s.c....n.DWs.Q.DVTKMSvIMI.MFL.AWSPYSIVCLWasFGDPk.Ipp.mAIiAPLFAKSSTFYNPCIYV.ANKkFRrA..aM..Cqt.q.......lPM......l.....
opsins ...Sf........Fi.PL.V.F.CY.......K....................VT.M.iiMI..FL..W.PY..V.............P....I...FAK.S..YNP.IYV..NK.FR........C..........................
.................................x.y....................................................................................................y...x...................

Curated Set of 48 vertebrate Peropsin Opsins

The sequences below represent all that are available as of March 2008. A few sequences such as chondrichthyes are incomplete. Note the main reference sequence collection contains additional deuterostome and even some lophotrochozoan peropsins.

>homSap Homo sapiens (human)
0 MLRNNLGNSSDSKNEDGSVFSQTEHNIVATYLIMA 1
2 GMISIISNIIVLGIFIKYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQ 0
0 VYAGLNIFFGMASIGLLTVVAVDRYLTICLPDV 1
2 GRRMTTNTYIGLILGAWINGLFWALMPIIGWASYAPDPTGATCTINWRKNDR 2
1 SFVSYTMTVIAINFIVPLTVMFYCYYHVTLSIKHHTTSDCTESLNRDWSDQIDVTK 0
0 MSVIMICMFLVAWSPYSIVCLWASFGDPKKIPPPMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRRAMLAMFKCQTHQTMPVTSILPMDVSQNPLASGRI* 0

>panTro Pan troglodytes (chimp)
0 MLRNNLGNSSDSKNEDGSVFSQTEHNIVATYLIMA 1
2 GMISIISNIIVLGIFIKYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQ 0
0 VYAGLNIFFGMASIGLLTVVAVDRYLTICLPDV 1
2 GRRMTTNTYIGLILGAWINGLFWALMPIIGWASYAPDPTGATCTINWRKNDR 2
1 SFVSYTMTVIAINFIVPLTVMFYCYYHVTLSIKHHTTSDCTESLNRDWSDQIDVTK 0
0 MSVIMICMFLVAWSPYSIVCLWASFGDPKKIPPPMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRRAMLAMFKCQTHQTMPVTSILPMDVSQNPLASGRI* 0

>gorGor Gorilla gorilla (gorilla)
0 MLRNNLGNGSDSKNEDGSVFSQTEHNIVATYLIMA 1
2 GMISIISNIIVLGIFIKYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQ 0
0 VYAGLNIFFGMASIGLLTVVAVDRYLTICLPDV 1
2 GRRMTTNTYIGLILGAWINGLLWALMPIIGWASYAPDPTGATCTINWRKNDR 2
1 SFVSYTVTVIAINFIVPLTVMFYCYYHVTLSIKHHTTSDCTESLNRDWSDQIDVTK 0
0 2
1 FRRAMLAMFKCQTHQTMPVTSILPMDVSQNPLASGRI* 0

>ponPyg Pongo pygmaeus (orang_abelii)
0 MLRNNLGNSSDSKNEDGSVFSQTEHNIVATYLIMA 1
2 GIISIISNIIVLGIFIKYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQ 0
0 VYAGLNIFFGMASIGLLTVVAVDRYLTICLPDI 1
2 GRRMTTNTYIGLILGAWINGLFWALMPIIGWASYAPDPTGATCTINWRKNDR 2
1 SFVSYTMTVIAINFIVPLTVMFYCYYHVTLSIKHHTTSDCTESLNRDWSDQIDVTK 0
0 MSVIMICMFLVAWSPYSIVCLWASFGDPKKIPPPMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRRAMLAMFKCQTHQTMPVTSILPMDVSQNPLASGRI* 0

>nomLeu Nomascus leucogenys (gibbon)
0 MLRNNLGNSSDSKNEDGSVFSQTEHNIVATYLIMA 1
2 GMISIISNIIVLGIFIKYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQ 0
0 VYAGLNIFFGMASIGLLTVVAVDRYLTICLPDV 1
2 GRRMTTNTYIGLILGAWINGLFWALMPIIGWASYAPDPTGATCTINWRKNDR 2
1 SFVSYTMTVIAINFIVPLTVMFYCYYHVTLSIKYHTTSDCTESLNRDWSDQIDVTK 0
0 MSVIMICMFLVAWSPYSIVCLWASFGDPKKIPPPMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRKAMLAMFKWPNHQTMPGTSILPMDVSQNPLTSGKI* 0

>macMul Macaca mulatta (rhesus)
0 MLRNNLGNSSDSKNEDGSVFSQTEHNIVATYLIMA 1
2 GMISILSNIIVLGIFIKYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQ 0
0 VYAGLNIFFGMASIGLLTVVAVDRYLTICLPDI 1
2 GRRMTTNTYIGMILGAWINGLFWALMPIIGWASYAPDPTGATCTINWRKNDR 2
1 SFVSYTMTVIAINFIVPLTVMFYCYYHVTLSIKHHATSDCTESLNREWSDQIDVTK 0
0 MSVIMICMFLVAWSPYSIVCLWASFGDPKKIPPPMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRRAMLAMFKCQTHQTMPVTSILPMDVSQNPLASGRI* 0

>papHam Papio hamadryas (baboon)
0 MLRNNLGNSSDSKNEDGSVFSQTEHNIVATYLIMA 1
2 GMISILSNIIVLGIFIKYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQ 0
0 VYAGLNIFFGMASIGLLTVVAVDRYLTICLPDI 1
2 GRRMTTNTYIGMILGAWINGLFWALMPIIGWASYAPDPTGATCTINWRKNDR 2
1 SFVSYTMTVIAINFIVPLTVMFYCYYHVTLSIKHHATSDCTESLNREWSDQIDVTK 0
0 MSVIMICMFLVAWSPYSIVCLWASFGDPKKIPPPMAIIAPLFAKSSTFYNPCIYMVANKK 2
1 FRRAMLAMFKCQTHQTMPVTSILPMDVSQNPLASGRI* 0

>calJac Callithrix jacchus (marmoset)
0 MLRNNLGNSSDSKNEDGSVFSQTEHNIVASYLIMA 1
2 GMISILSNIIVLGIFIKYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQ 0
0 VYAGLNIFFGMASIGLLTVVAADRYLTICLPDI 1
2 GRRMTTSTYIIMILGAWINGLFWALMPIVGWASYAPDPTGATCTINWRKNDR 2
1 SFVSYTMAVIAINFIVPLTVMFYCYYHVTLFIKHHTTSDCTESLNRDWSDQIDVTK 0
0 MSVIMICMFLVAWSPYSIVCLWASFGDPKNIPPAMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRRAMLAMLKCQTHQTMPVTSVLPMDISQNPLASGRI* 0

>tarSyr Tarsius syrichta (tarsier)
0 MLKNDLGNSSDSKNEDGSVFSQTEHNIVATYLIMA 1
2 GMISILSNIIVLGIFVKYKELRTPTNAIIINLSVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQ 0
0 IYAGLNIFFGMASIGLLTVVAVDRYLTICRPDI 1
2 GRRMTTNTYVGMILGAWINGLFWASMPIAGWATYAPDPTGATCTINWRKNDA 2
1 TFVSYTMAVIAINFIVPLTVMFYCYYHATQSIKHHVASNCVESLNRDWSDQVDVTK 0
0 2
1 FRRAMFAMLKCQTYQAMPATSSLPMNVSQNPLTSGKN* 0

>otoGar Otolemur garnettii (bushbaby)
0 MLRNNLGNNSDSKNEDGSVFSQTEHNIVATYLIMA 1
2 GMISILSNIIVLGIFIKYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLHGSWKFGYAGCQ 0
0 IYAGLNIFFGMASIGLLTVVAVDRYLTICRPDI 1
2 GRRMTTNSYIGMILGAWVNGLFWALMPITGWASYAPDPTGATCTINWRKNDA 2
1 SFVSYTMTVIAVNFVVPLMVMFYCYYHVTQSIKRHTATNCTESLNRDWSDQVDVTK 0
0 MSVIMICMFLVAWSPYSIVCLWALFGDPKKIPPSMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRRAMFAMFKCQTHQAMAVTSILPMDISQNPLASRRI* 0

>micMur Microcebus murinus (mouse_lemur)
0 1
2 0
0 IYAGLNIFFGMASIGLLTVVAMDRYLTICRPDI 1
2 GRRMTTHTYVGMILGAWVNGLFWAVMPITGWAGYAPDPTGATCTINWRKNDA 2
1 0
0 MSVIMICMFLVAWSPYAIVCLWASFGNPEKIPPSMAIIAPLFAKSSTFYNPCIYVIANKK 2
1 FRRAMFAMFKCQTHQAMPVTSIFPMGVSQNPLPSGRT* 0

>tupBel Tupaia belangeri (treeshrew)
0 MLGSDAGNSSNLGNEDSSAFSQTEHNIVAAYLLTA 1
2 GVISILSNIVVLGIFVTHKELRTPTNAIIINLAFTDIGVSSIGYPMSAASDLHGRWKFGHAGCQ 0
0 IYAGLNIFFGMASIGLLTVVAVDRYLTLCRPAV 1
2 GRRMGSSTYAAMILGAWLNGLFWAAMPIAGWAGYAPDPTGATCTINWRKNDA 2
1 SFVSYTMAVIAVNFVVPLTVMSYCYVLVARAIAGHAASSCSEHRCRDWSEQVDVTK 0
0 MSVLMILMFLVAWSPYSIVCLWASFGDPQRIPPAMAIVAPLFAKSSTFYNPCIYVLANKK 2
1 FRKAMCAMFKCQTHQAMSVTSVLPMASSPRPLAPARV* 0

>musMus Mus musculus (mouse)
0 MLSEASDNSSGSRSE-GSVFSRTEHSVIAAYLIVA 1
2 GITSILSNVVVLGIFIKYKELRTPTNAVIINLAFTDIGVSSIGYPMSAASDLHGSWKFGHAGCQ 0
0 IYAGLNIFFGMVSIGLLTVVAMDRYLTISCPDV 1
2 GRRMTTNTYLSMILGAWINGLFWALMPIIGWASYAPDPTGATCTINWRNNDT 2
1 SFVSYTMMVIVVNFIVPLTVMFYCYYHVSRSLRLYAASDCTAHLHRDWADQADVTK 0
0 MSVIMILMFLLAWSPYSIVCLWACFGNPKKIPPSMAIIAPLFAKSSTFYNPCIYVAAHKK 2
1 FRKAMLAMFKCQPHLAVPEPSTLPMDMPQSSLAPVRI* 0

>ratNor Rattus norvegicus (rat)
0 MLRDALDNSSGSGSE-GSVFTKSEHSIIAAYLIVA 1
2 GIISILSNIIVLGIFIKYKELRTPTNAVIINLAFTDIGVSSIGYPMSAASDLHGSWKFGHAGCQ 0
0 VYAGLNIFFGMVSIGLLTVVALDRYLTISCPDV 1
2 GRRMTGNTYLSMVLGAWINGLFWALMPIVGWASYAPDPTGATCTINWRKNDT 2
1 SFVSYTMMVIVVNFIVPLTVMFYCYYHVSQSMRLSAASNCTTHLNRDWAHQADVTK 0
0 MSVMMILMFLLAWSPYSVVCLWACFGNPKKIPPSLAIIAPLFAKSSTFYNPCIYVAANKK 2
1 FRKAMFAMLKCQPHQAMPEPSTLAMGVPHSPLAPARI* 0

>speTri Spermophilus tridecemlineatus (ground_squirrel)
0 1
2 0
0 1
2 2
1 SFVSYTMTVIAVNFIVPLTVMFYCYYHVTRSIKHHAASNCTAYLNRDWSDQVDVTK 0
0 MSVIMILMFLVAWSPYSIVCLWASFGNPKKIPPSMAIIAPLFAKSSTFYNPCIYVAANKR 2
1 FRRAMFAMFKCQTHQAMPVTSVLPMDVSQSPRASGRI* 0

>dipOrd Dipodomys ordii (kangaroo_rat)
0 MLRNNVGNSSGSRNEDGSVFSQTEHNIVATYLITA 1
2 GVISILSNLIVLGIFIKYKELRTPTNAIIINLALTDIGVSSIGYPMSAASDLYGRWKFGYAGCQ 0
0 IYAGLNIFFGMASIGLLTVVAIDRYLTICHPDI 1
2 GRGMTTRTYVTMILGAWINGLFWALMPIIGWASYAPDPTGATCTINWRKNDT 2
1 0
0 MSVVMILMFLVAWSPYSIVCLWASFGDPKEIPPPMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRRAMLAMLKCQTHQAMPVTSILPMDVSQNPLASGRI* 0

>cavPor Cavia porcellus (guinea_pig)
0 MLRHSLGNSSDSKNEDGSVFSQTEHNIVAAYLILA 1
2 GLISILSNIIVLGIFIKYKELRTPTNAIIMNLALTDIGVSSIGYPMSAASDLHGSWKFGYAGCQ 0
0 VYAGLNIFFGMASIGLLTVVAVDRYLTICRPDI 1
2 GRRMTSHSYVGMILGAWINGLFWALMPIIGWASYAPDPTGATCTINWRKNDV 2
1 AFVSYTMTVIAINFIVPLAVMFYCYLHITRAIRRHVAGDRPPNLSGDWSDQVDVTK 0
0 MSVVMILMFLVAWSPYSIVCLWASFGDPRRISPSMAIIAPLFAKSSTFYNPCIYVIANKK 2
1 FRRAMFAMFQCQTHQAVPVASILPMDASQSPLASGRI* 0

>oryCun Oryctolagus cuniculus (rabbit)
0 MLRNNLSNSSDFKHEDGSVFSQTEHNIVATYLILA 1
2 GMISILSNLIVLGIFIKYKELRTPTNAIIINLAFTDIGVSSIGYPMSAASDLHGSWKFGYAGCQ 0
0 IYAGLNIFFGMASIGLLTVVAMDRYLTICHPDV 1
2 GRRMTTRTYLGLILGAWVNGLFWALMPIAGWASYAPDPTGATCTINWRKNDT 2
1 SFVSFTMAVIAINFVVPLTVMFYCYYHVTQSIKQHRASDCTEYLNRDWSDQVDVTK 0
0 MSVIMIFMFLVAWSPYSIVCLWASFGDPKKIPPAMAIIAPLFAKSSTFYNPCIYVAANKR 2
1 FRRAMFAMFKCQTHQAMPVTSVLPMDVSQNPLPSGII* 0

>ochPri Ochotona princeps (pika)
0 MLRHNLGNSSEAKVEAGSVFSQTEHNIVAAYLILA 1
2 GMISILSNLIVLGIFIKYKELRTPTNAIIINLAFTDIGVSSIGYPMSAASDLHGSWKFGYAGCQ 0
0 IYAGLNIFFGMASIGLLTVVAMDRYLTICQPDI 1
2 GRRMTTHTYFGMILGAWINGLFWALMPIVGWASYAPDPTGATCTINWRKNDK 2
1 SFVSFTMAVIMVNFVVPLTVMFYCYYYVTQSIKHHTASDCTKSLNRDWSDQVDVTK 0
0 MSVIMILMFLVAWSPYSIVCLWASFGDPQKIPPSMAIIAPLFAKSSTFYNPCIYVAANKR 2
1 SRRAMFAMFKCQIPQAKPVTSLSPRDVSQSPLSSGRT* 0

>canFam Canis familiaris (dog)
0 MLRNNLDNSTDSKNEDGSVFSQTEHNIVAAYLITA 1
2 GIISIFSNLIVLGIFIKYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQ 0
0 IYAGLNIFFGMASIGLLTVVAMDRYLTICSPDT 1
2 GRRMTTNTYISMILGAWLNGLFWALMPIIGWASYAPDPTGATCTINWRKNDA 2
1 FFVSFTMTVIAVNFIVPLTVMFYCYYHVTRSIKCHTTSNCTEYLNRDWSDQVDVTK 0
0 MSVIMIFMFLVAWSPYSIVCLWASFGDPKKIPPSMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRKAIFAMFKCQTHQAMPGTSILPMDVSQNPLASGRN* 0

>felCat Felis catus (cat)
0 MLRNNLDNSSDSKNEDGSVFSQTEHNIVAAYLITA 1
2 0
0 IYAGLNIFFGMASIGLLTVVAMDRYLTICSPNS 1
2 GRRMTTNTYISMILGAWLNGLFWALMPIIGWASYAPDPTGATCTINWRKNDA 2
1 SFVSFTMTVIAINFNVPLTVMFYCYYHVTQTIKCHDTSNCTGYLNRDWSNQVDVTK 0
0 2
1 FRKAMFAMFKCENRQPMPVTSILPMDVSQNPLTSGRK* 0

>bosTau Bos taurus (cow)
0 MLRNNLGNSSDCKNENGSVFSQTEHNIVAAYLITA 1
2 GVISILSNIIVLGIFIKFKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLHGSWKFGYAGCQ 0
0 IYAGLNIFFGMASIGLLTVVAVDRYLTICHPDA 1
2 GRRMTANTYISMILGAWTNGLFWALMPIIGWASYAPDPTGATCTINWRKNDV 2
1 SFVSYTMMVVAINFIVPLTVMFYCYYHVTQSIKHHGTNNCTEYLNRDWSDQVDVTK 0
0 MSVIMILMFLVAWSPYSIVCLWASFGDPKKIPPSMAIIAPLFAKSSTFYNPCIYVIANKK 2
1 FRRAMLAMFKCQTTQAMPVTSVLPMDVPQNPLTSGKV* 0

>turTru Tursiops truncatus (dolphin)
0 MLRNNLGNSSDSKNEDGSTFSQTEHNIVAAYLITA 1
2 GMISVLSNIIVLGIFIKYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLHGSWKFGYAGCQ 0
0 IYAGLNIFFGMASIGLLTVVAVDRYLTICCPGA 1
2 GRRMTTNTYISMILGAWFNGLFWALMPIVGWASYAPDPTGATCTINWRKNDV 2
1 SFVSYTMMVVAINFIVPLIMMFYCYYHVTRSIKHHATSNCTEYLNRDWSDQVDVTK 0
0 MSVIMILMFLVAWSPYSIVCLWASFGDPKKIPPSVAILAPLFAKSSTFYNPCIYVIANKK 2
1 FRRAMLAMFKCQTHQAMPMESILPMDVPQNPLTSGKV* 0

>susScr Sus scrofa (pig)
0 MLMDLGNNSDSKNEDGSVFSQTEHSIVAAYLITA 1
2 GMISILSNIVVLGIFIKHKELRTPTNAIIINLAATDIGVSSIGYPMSAASDLHGSWKFGYAGCQ 0
0 IYAGLNIFFGMASIGLLTVVAVDRYLTICRPEA 1
2 GRRMTTNTYISMILGAWINGLFWALMPIIGWASYAPDPTGATCTINWRKNDV 2
1 SFVSYTMTVVAVNFIVPLIVMFYCYYHVTQSIKSHATSNCTAYLNRDWSDQVDVTK 0
0 MSVIMILMFLVAWSPYSIVCLWASFGDPKKIPPPMAIIAPLFAKSSTFYNPCIYVIANKK 2
1 FRRAMLAMFKCQTHQAMPLESTLPMDVPQNPLASGRV* 0

>vicVic Vicugna vicugna (vicugna)
0 MLRNNLGNSSDSKNEYGSVFSQAEHNIVAAYLITA 1
2 LRTPTNAIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQ 0
0 IYAGLNIFFGMASIGLLTVVAMDRYLTICRPDA 1
2 GRRMTTNTYISMILGAWINGLFWASVPIIGWAGYAPDPTGATCTINWRQNDV 2
1 SFVSYTMMVVAINFIVPLIVMFYCYYHVTRSIKCRATSNCTEYLNRDWSDQVDVTK 0
0 MSVIMILMFLLAWSPYSIVCLWASFGDPKKIPPSMAIIAPLFAKSSTFYNPCIYVIANKK 2
1 FRRAMLAMFKCQTHQAMPMTSILPMDVPQNPLTSGRL* 0

>equCab Equus caballus (horse)
0 MLRNNLGNSSDSKNEDGSVFSQTEHNIVAAYLIMA 1
2 GMISILSNIIVLGIFVKYKELRTSTNAIIINLAVTDIGVSSIGYPMSAASDLYGRWKFGYAGCQ 0
0 IYAGLNIFFGMASIGLLTVVALDRYLTTCRPDA 1
2 GRRMTTSTYTSMILGAWVNGLFWALMPIVGWANYAPDPTGATCTINWRKNDA 2
1 SFVSYTMAVVAINFIVPLTVMFYCYYHVTRSMKRHPTSNCTQYLNTDWSDQVDVTK 0
0 MSVIMIFMFLVAWSPYSIVCLWASFGDPKKIPPSMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRRAMFAMFKCQTHRAMPVTSILPMDVPQNQLASGRI* 0

>myoLuc Myotis lucifugus (microbat)
0 MLKNNLSNSSDFKNEDGSVFSQTEHNIVATYLIMA 1
2 GMISILSNVIVLGIFITYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLHGSWKFGYAGCQ 0
0 1
2 GRKMTTHTYISMILGAWINGLFWALMPIIGWASYAPDPTGATCTINWRKNDV 2
1 SFVSFTMTVIAINFIVPLTVMFYCYYHVTRSIKCRATSNCTEYLNRDWADQVDVTK 0
0 MSVIMIFMFLVAWSPYSIVCLWASFGDPKKIPPSMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRRAMFAMFKCQTHQTMTTMSFLPMDVPQNPLTSGRI* 0

>pteVam Pteropus vampyrus (macrobat)
0 MLRNNLGNSSNSKNEDGSVFSQTEHNIVATYLIIA 1
2 GMISILSNIIVLGIFFKYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQ 0
0 1
2 GRRMTTNTYIIMILGAWINGLFWALMPIIGWASYAPDPTGATCTINWRKNDA 2
1 SFVSFTMTVIAINFIVPLAVMFYCYYHVTRSIKCHTTSNCTEYLNRDWSDQVDVTK 0
0 MSVIMIFMFLVAWSPYSIVCLWASFGDPKKIPPSMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRRAMFAMFKCQDHQSMPVTSVLPMDVPQNPLTSGRI* 0

>sorAra Sorex araneus (shrew)
0 MLENRSDNSSGPTPEGGAVLSRTEHRLVAAYLIVA 1
2 GVISILSNTAVLGIFIKYKELRTRTNAIIMNLALTDIGVSSIGYPMSAASDLHGSWRFGHAGCQ 0
0 VYAGLNIFFGMASIGLLTLVAVDRYLTLCRPDA 1
2 GRSMTTNSYVGLILGAWINGFFWALMPILGWASYAPDPMGATCTINWRGNDV 2
1 SFVSYTMTVVAVNFVMPLTVMAYCYYHVIRSL-----HSSPGHLDGDWSSQVDVTK 0
0 MSVVMILAFLAAWSPYSIVCFWASFGDPKKIPPSMAVIAPLFAKSSTFYNPCIYVVANKK 2
1 FRRAMSAMLTCRAQGAMPAASTLPMDAAHSPQASGRN* 0

>eriEur Erinaceus europaeus (hedgehog)
0 MLENNLGNGSDSENEDGSVFSQTEHNIVAAYLIVA 1
2 0
0 IYAGLNIFFGMASIGLLTVVAVDRYLTICRPHT 1
2 GRSMSANSYIAMILGAWLNGLFWALMPISGWAGYAPDPTGATCTINWRENDG 2
1 SFVSYTMTVIAINFMVPLAVMLYCYYHVTRSMKCHTARSCAEYVHGDWSDQVDVTK 0
0 MSVIMILMFLVAWSPYSVVCLWASFGDPKKIPPSMAIMAPLFAKSSTFYNPCIYVLANKK 2
1 FRRAMFAMFKCQTHQAMPVTNTLPMDIPQK* 0LDSRRN* 0

>loxAfr Loxodonta africana (elephant)
0 MLRNSLDNSSDSKNEDASVFSQTEHNIVATYLIMA 1
2 GMISILSNIIVLGIFIKYKELRTPTNAIIINLAVTDIGVSSIGYPMSAASDLHGRWKFGYTGCQ 0
0 IYAGLNIFFGMASIGLLTVVAVDRYLTICHPHI 1
2 GRRMTSNTYVSMILGAWINGLLWALLPITGWASYAPDPTGATCTINWRKNDA 2
1 SFVSYTMTVIVINFVVPLAVMFYCYYHVTRSIKRHTASNCAEYLNRDWSDQLDVTK 0
0 MSVIMILMFLVAWSPYSIVCLWASFGDSKKIPPSMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRRAMFAMFKCQTHQAEPVTCILPMNVSQNPLAAGRI* 0

>proCap Procavia capensis (hyrax)
0 MLRNNLGNSSDSKNEEGSVFSQTEHNIVAAYLIMA 1
2 0
0 IYAGLNIFFGMSSIGLLTVVAIDRYLTICHPNV 1
2 2
1 SFVSYTVTVIMVNFVVPLAVMFYCYYHITLSIKCHAARNCAEPLCRDWSDQLDVTK 0
0 MSVIMILMFLMAWSPYSIVCLWASFGDPTKIPPSMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRRAMLAMFKCQTHQAVPVTNILPMTVSQNSSASGRI* 0

>echTel Echinops telfairi (tenrec)
0 MLKNNEGTSSGSHSEEGSVFSQAEHRIVASYLITA 1
2 GMISILSNFIVLGIFIKYKELRTPTNAMIINLAVTDIGVSSIGYPMSAASDLHGSWKFGYAGCQ 0
0 VYAGLNIFFGMASIGLLTAVAVDRYLTICHPDR 1
2 GRRMTSNTYVGMILGAWINGFLWALLPVIGWASYAPDPTGATCTINWRENDA 2
1 SFVSYTMTVVGINFVVPLAVMFYCYYHVTQTIKRQTASNGAENLHRDWSDQLDVTK 0
0 2
1 FRRAMFALLQCQPQEARRVTSILPMNVSQNPMASGRL* 0

>dasNov Dasypus novemcinctus (armadillo)
0 MLRNNTGNSSDSKDEDGSVFSQTEHNIVATYLIMA 1
2 GIISIFSNIIVLSIFIKYKELRTPTNTIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQ 0
0 IYAGLNIFFGMTSIGLLTVVAMDRYLTICRPDT 1
2 GRRMTINTYISMILGAWINGLFWALMPIAGWASYAPDPTGATCTINWRKNNI 2
1 SFVSYTMTVVAINFLVPLAVMLYCYYHVTRSIKHHTKSNCSEYFNRDWSNQVDVTK 0
0 MSVIMISMFLVAWSPYSIVCLWASFGDPEKIPPSMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRRAIFAMLKCQTHQAMPVMSILPMNVSENPLASGRI* 0

>choHof Choloepus hoffmanni (sloth)
0 1
2 GIISILSNIVVLGIFIKYKELRTPTNTVIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQ 0
0 IYAGLNIFFGMASIGLLTVVAVDRYLTICHPDV 1
2 GRRMTINTYISMILGAWINGLFWALLPVTGWASYAPDPTGATCTINWRKNDG 2
1 SFVSYTMTVVVINFVVPVAVMFYCYYHITRSIKRRNSSNCSEYLNRDWSDQVDVTK 0
0 MSVIMILMFLLAWSPYSIVCLWASFGDPKKIPPSMAIIAPLFAKSSTFYNPCIYVVANKK 2
1 FRTIMFAMLKCQTHQAVPVTSILPMNVSENPLASGRI* 0

>macEug Macropus eugenii (wallaby)
0 MFQNDSLEPEKESYSVFSPTEHNIVAAYLITA 1
2 GVISIPSNIIVLGIFVKYKELRTATNTIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYAGCQ 0
0 IYAGLNIFFGMASIGLLTAVAIDRYLTICQPDL 1
2 2
1 SFVSYTMTVIAINFVMPLVVMFYCYYNVSLKMKQYTRSSCPEHINRDWSNQVDVTK 0
0 MSVIMILMFLLAWSPYSVVCLWASFGDPKEIPPAMAIIAPLFAKSSTFYNPCIYVAANKK 2
1 FRRAISAMMRCETHQSMPVSNALPLNLT* 0

>monDom Monodelphis domestica (opossum)
0 MFKNNSVKTLAPEKEGPSVFSPIEHKIVAAYLITA 1
2 GVISIVSNVIVLGIFVKYKALRTATNTIIINLAVTDIGVSSIGYPMSAASDLYGSWKFGYDGCQ 0
0 IYAGLNIFFGMASIGLLTAVAIDRYLTICQPDL 1
2 GGRMTSYNYTLMILTAWVNGFFWALMPIVGWAGYAPDPTGATCTINWRKNDV 2
1 SFVSYTMTVITINFAMPLGVMFYCYYNVSQKMKQYSPSNCPDHINRDWSNQVAVTK 0
0 MSVVMILMFLLAWSPYSIVCLWASFGDPKEIPPAMAIVAPLFAKSSTFYNPCIYVAANKK 2
1 FRRAISAMIRCQTHQSMPISNALPMN* 0

>ornAna Ornithorhynchus anatinus (platypus)
0 MRRNDSANLLESEHHDRSAFSQTDHNIVAAYLITA 1
2 GIMSIVSNVIVLGIFVKFEELRTATNAIIINLAVTDIGVSGIGYPMSAASDLHGSWKFGHAGCQ 0
0 IYAGLNIFFGMSSIGLLTVVAVDRYLTICRPAI 1
2 GRKMTRSNYTAMILAAWMNGFFWASMPLLGWASYASDPTGATCTINWRKNDA 2
1 SFISYTMTVIAVNFAVPLIVMFYCYYNVSKAMRQYPASRVLENLNIDWSEQVDVTK 0
0 MSVVMILMFLMAWSPYSIVCLWSSFGDPKKISPAVAIMAPLFAKSSTFYNPCIYVVANKK 2
1 FRRAMLSMVQCQTHREITITDVLPMNRSRSPLTL* 0

>galGal Gallus gallus (chicken)
0 MHWNDSANSSESDAEAHSVFTQTEHNIVAAYLITA 1
2 GVISIFSNIVVLGIFVKYKELRTATNAIIINLAFTDIGVSGIGYPMSAASDLHGSWKFGYTGCQ 0
0 IYAALNIFFGMASIGLLTVVAVDRYLTICRPDI 1
2 GRRMTTRNYAALILAAWINAVFWASMPTVGWAGYASDPTGATCTANWRKNDV 2
1 SFVSYTMSVIAVNFVVPLTVMFYCYYNVSRTMKQYTSSNCLESINMDWSDQVDVTK 0
0 MSVVMIVMFLVAWSPYSIVCLWSSFGDPKKISPAMAIIAPLFAKSSTFYNPCIYVIANKK 2
1 FRRAILAMVRCQTRQEITISNALPMTVSLSALTS* 0

>taeGut Taeniopygia guttata (finch)
0 MHWNDSSNSSESDDEAHSAFTQTEHNIVAAYLITA 1
2 GVISIFSNIVVLGIFVKYKELRTATNAIIINLAFTDIGVSGIGYPMSAASDLHGSWKFGYTGCQ 0
0 IYAALNIFFGMASIGLLTVVAVDRYLTICRPDI 1
2 GRRMTTRSYATLILAAWINAVFWSSMPTAGWASYAPDPTGATCTVNWRKNDA 2
1 SFISYTMSVIAVNFVVPLTVMFYCYYNVSRTMKQYASSNCLESINIDWSDQVDVTK 0
0 MSVVMIIMFLVAWSPYSIVCLWSSFGDPKKISPAMAIIAPLFAKSSTFYNPCIYVIANKK 2
1 FRRAILAMVRCQTRQEITINNALPMSVSQSALTSQNSSHLPA* 0

>anoCar Anolis carolinensis (lizard)
0 MFLNDSANSSESDDEPHSAFSQAEHNIVAAYLITA 1
2 GVISLLSNIVVLGIFVKYKELRTATNAIIINLAFTDIGVSGIGYPMSAASDLHGSWKFGYTGCQ 0
0 IYAALNIFFGMASIGLLTVVAIDRYLTICKPHI 1
2 GSRLTATNYTTLILAAWINALFWASMPVVGWASYAPDPTGATCTVNWRKNDT 2
1 SFVSYTMSVIAVNFVIPLSVMFYCYYNVSKTMKYYMRNSCLENINIDWSDQVDVTK 0
0 MSVVMIIMFLLAWSPYSIVCLWSSFGDPKKISPAMAIVAPLFAKSSTFYNPCIYVIANKR 2
1 FRRAILAMIRCQTRQEITINNVLPMSVSQSTIA* 0

>xenTro Xenopus tropicalis (frog)
0 METLAEVSTLLPAGTGTVNISDASSEVHSVFSQSEHNIVAAYLITA 1
2 GVISILSNIIVLGIFVKYKELRTATNAIIINLAFTDIGVSGIGYPMSAASDLHGSWKFGYVGCQ 0
0 IYAGLNIFFGMASIGLLTVVAIDRYLTICRPDI 1
2 GRRISGRHYTAMILAAWINAVFWSVMPVVGWSSYAPDPTGATCTINWRKNDV 2
1 SFVSYTMSVVAVNFVVPLMVMFYCYYNVSRTMKGYGSRSSLGGINADWSDQTDVTK 0
0 MSMVMIVMFLVAWSPYSIVCLWSSFGDPRKIPPAMAIIAPLFAKSSTFYNPCIYVIANKK 2
1 FRRAILSMVQCKSRQEVTLDNHFPMNVSQSTLTT* 0

>danRer Danio rerio (zebrafish)
0 MESGLLNVSAETVYGEKSAFTQTEHNIVAAYLITA 1
2 GVISLSSNIVVLLMFVKFRELRTATNAIIINLAFTDIGVAGIGYPMSAASDLHGSWKFGYMGCQ 0
0 IYAALNIFFGMASIGLLTVVAIDRYLTICRPDI 1
2 GQKLTTRSYTLLIVAAWLNAVFWSSMPIVGWAGYAPDPTGATCTINWRNNDT 2
1 SFVSYTMTVITVNFIIPLSVMFYCYYNVSATVKRFKASNCLDSINMDWSDQMDVTK 0
0 MSVIMIVMFLAAWSPYSIVCLWASFGDPQKIPAPMAIIAPLLAKSSTFYNPCIYVIANKK 2
1 FRRAIIGMIRCQTRQRVTINNQLPMMASSVPLNP* 0

>takRub Takifugu rubripes (fugu)
0 MKVFSLVNISNEVEGKSVFTQWEHNIVAGYLIIA 1
2 GVISLTSNIVVLLMFVKFKELRTATNFIIINLAFTDIGVAGIGYPMSAASDIHGSWKFGHTGCQ 0
0 IYAALNIFFGMASIGLLTVVAIDRYITICRPDI 1
2 GRKMTVQSYNLLILAAWLNAVFWSSMPVVGWAAYAPDPTGATCTINWRQNNA 2
1 SFISYTMAVIGVNFMLPLFVMFYCYYNVSVTVNRCKTSNCLDDINIEWSEQMDVTK 0
0 MSLVMIIMFLVAWSPYSIVCLWASFGDPKAIPAPMAIIAPLFAKSSTFYNPCIYVIANKK 2
1 FRRAIIGMIRCQTRQQMTINTEIPMTTSQQTATQ* 0

>tetNig Tetraodon nigroviridis (pufferfish)
0 MKSVNSSNDVEGKSAFTQWEHNIVAGYLITA 1
2 GIISLTSNIVVLLMFVKFKELRTATNFIIINLAFTDIGVAGIGYPMSAASDIHGSWKFGHTGCQ 0
0 IYAALNIFFGMASIGLLTVVAIDRYLTICRPDI 1
2 GRKMTVQSYNLLIAAAWLNAVFWSSMPVVGWAAYAPDPTGATCTINWRQNNV 2
1 SFISYTMAVISVNFILPLFVMFYCYYNVSVTVKQYKANNCLDNINIEWSEQMDVTK 0
0 MSIVMIIMFLVAWSPYSIVCLWASFGDPKTISAPMAIIAPLFAKSSTFYNPCIYVITNKK 2
1 FRQAIIGMIRCQTRQQITINTDIPMTASQQTLTQ* 0

>gasAcu Gasterosteus aculeatus (stickleback)
0 MGIDPEVNVTDDVTLYGGKSAFTQLEHNIVAGYLITA 1
2 GVISLFSNIVVLLMFWKFKELRTATNFIIINLAFTDIGVAGIGYPMSAASDIHGSWKFGYAGCQ 0
0 IYAALNIFFGMASIGLLTVVAIDRYLTICRPDI 1
2 GQKMTMQSYNLLILAAWLNAVFWSSMPVVGWASYAPDPTGATCTINWRQNDV 2
1 SFISYTMAVIAVNFVLPLSAMFYCYYNVSATVKRYKASNCLDSANIDWSDQMDVTK 0
0 MSIVMIIMFLVAWSPYSIVCLWASFGDPKTIPAPMAIIAPLFAKSSTFYNPCIYVIANKK 2
1 FRRAIIGMVRCQTRQRITINSQVPMTTSQQPLTQ* 0

>oryLat Oryzias latipes (medaka)
0 DSAVNVSDAAAPYGGKSAFTQLEHNIVAGYLITA 1
2 GVISLSSNVLVLLMFVKFRELRTATNIIIINLAFTDVGVAGIGYPMSAASDIHGSWKFGYTGCQ 0
0 IYAALNIFFGMASIGLLTVVAIDRYLTICRPDL 1
2 GQKMTMQSYNLLILAAWLNAVFWSSMPIVGWAGYAPDPTGATCTINWRQNDA 2
1 SFVSYTMAVIAVNFVVPLTVMFYCYYNVSATVKRYKASNCLDSINIDWSDQMDVTK 0
0 MSIVMIVMFLVAWSPYSMVCLWASFGDPKTIPAPMAIIAPLFAKSSTFYNPCIYVIANKK 2
1 FRRAIIGMIRCQTRQRITISTQVPMTISQQPLTQ* 0

>calMil Callorhinchus milii (elephantfish)
0 1
2 0
0 1
2 2
1 LFVSYTMTVIAVNFVVPLSVMFFCYYNVSKTMSRFISSPSPENINLDWSDQLDVTK 0
0 MSVVMIVMFLLAWSPYSIVCLWASFGNPKLIPPAMAIIAPLFAKSSTFYNPCIYVIANKK 2
1 FRKAIMAMICCQNRQEITINHTLPMTISRVPLTE* 0