Opsin evolution: Neuropsin phyloSNPs
See also: Curated Sequences | Peropsins | RGR phyloSNPs | LWS | Encephalopsins | Melanopsins | Update Blog
Neuropsin backgrounder
Neuropsin (OPN5) is a deeply diverged member of the opsin family with a single publication on it and considerable confusion over the name (mostly used for an unrelated kalikrein serine protease KLK8, not an opsin). There are no known disease associations or described knockout phenotypes; it is expressed primarily in brain, spinal chord, and testes.
Neuropsin has all the classical attributes of a rhodopsin-class GPCR and indeed opsin photoreceptor: Schiff base lysine at expected position, standard tyrosine counterion and DRY motif, seven transmembrane configuration, disulfide at expected position, proximal glycosylation and distal palmitoylation and kinase sites. It is most closely related to peropsin and rgropsin in terms of blast clustering and intron positioning. Its G-protein signaling partner is not known.
Its evolution is illuminated by the massive comparative genomics study described here, which extracts and compares over 50 full length sequences from various genomics projects. Neuropsin can be located outside chordates but not outside deuterostomes. However, like peropsin and rgropsin, it must have originated much earlier in pre-Bilaterans. Thus its absence in earlier diverging species must be due either to gene loss or unrecognizability. Some role in deuterostomes however has persisted in deuterostomes over billions of years of branch length.
Within placental mammals, neuropsin is extraordinarily conserved, with percent identity relative to human protein 96% averaged over 31 species (likely above the 95% percentile of all coding genes proteomewide). That conservation drops considerably at marsupials and monotremes (86%), is less striking at tetrapods (78%), and not especially remarkable at teleost fish (68%). This pattern suggests neuropsin acquired significant new adaptive functionality on the placental mammal stem, leading to marked resilience to fixation of any further variation.
The structure of the neuropsin gene is rather odd at the 3' end. In human, a weak splice signal appears to have developed that results, after an intron of 12,244 bp, in a seventh very short exon followed by a long 3'UTR. However a stop codon is soon encountered if the splice is not taken (which it is in 6 of 7 transcripts). This results in two slightly different alternative carboxy termini sequences QEV* vs QEWE*. Very few transcripts exist in this region for any species but it appears that the recent ancestral form of the protein only utilized the initial stop codon in exon 6.
This feature GTatga is conserved in all species back to platypus; indeed an unexplained conservation in nucleotide sequence extends well beyond this. However the splice acceptor and WE* appear an option only back to rhesus, whereas the QEV* option is available in all 38 available mammal genomic sequences. The Q itself is a strong phyloSNP characteristic of mammals from platypus through lemurs.
A single mouse transcript also terminates early; four others continue on. No species other than rat has an available transcript in this region. No other rodent genomes could have an orthologous exon: kangaroo rat and ground squirrel have frame shifts, pika lacks the splice acceptor, and rabbit and guinea pig have no homologous sequence.
This can be explained by independent origins of splice acceptors in various clades. However far more comparative transcripts are needed to understand the 3' end of this gene. It may be that the conserved intronic sequence following exon 6 is an accident waiting to happen as it conserves -- for whatever reason -- half of a splice site.
Novel neuropsins in amphioxus and sea urchin
The genome of Branchiostoma (amphioxus, lancelet) contains two distinct neuropsins about 75% identical to each other and 42% to human. These cluster unambiguously with vertebrate opsins and share critical conserved residues. An extra intron distinguishes them from the vertebrate neuropsin pattern. Recall Branchiostoma species has three rather diverged (and well-studied) peropsins but no evident Rgr opsin. These raises the question whether neuropsin and peropsin developed substantial visual roles in this species as an alternative to the ciliary imaging opsin pathway seen by lamprey divergence. Sea urchins, but not acornworm Saccoglossus, contain a single neuropsin that is quite diverged.
These neuropsins are newly reported here, meaning they were not localized in recent in situ hybridization studies. That's especially unfortunate in view of the antecedent role the Branchiostoma ancestral node plays in the evolution of chordate eye and the complexities of photoreceptor tissues in the extant species.
PhyloSNPs in vertebrate neuropsins
Alignment analysis coming shortly. Neuropsin has rather few of them.
position ...................................................................................................1.........1.........1.........1.........1.........1.........1.........1......1.. position .........1.........2.........3.........4.........5.........6.........7.........8.........9.........0.........1.........2.........3.........4.........5.........6.........7......7.. position 123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567.. excMemCy eeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeMMMMMMMMMMMMMMMMMMMMMccccccccccccccccccccMMMMMMMMMMMMMMMMMMMMMeeeeeeeeeeeeeMMMMMMMMMMMMMMMMMMMMMcccccccccccccccccccccMMMMMMMMMMMMMMMMMMMMMeeeeee.. keyResid ...GLC.................................................................................................diS..cIon.................DRY............................................... exonNumb 111111111111111111111111111111111111111111122222222222222222222222222222222222222223333333333333333333333333333333333333333333333333333333334444444444444444444444444444444444444.. 10homSap MALNHTALPQDERLPHYLRDGDPFASKLSWEADLVAGFYLTIIGILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGISVVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSYGVWLKRKHAYICLAAIWAYASFWTTMPLVGLGDYVPE.. 11panTro ................................................................................................................................................................................... 12gorGor ............................................................................................................................................------------------..................... 13ponPyg ................................................................................................................................................................................... 14nomLeu ................................................................................................................................................................................... 15rheMac .................................I............................................................................................................................................A.... 16papHam .................................I............................................................................................................................................A.... 17calJac ......S.......................................................................................................................................................................A.... 18tarSyr ....................................................................................I.........................................................................................A.... 19otoGar ............LR..........................................................V...................................................................----------------------..M......E....... 20micMur ......V.................................I..........................................---------------------------------------------------------..............L....V..............A.... 21tupBel ............S.............................................................................................................................................V...................A.... 22musMus ....................E..................................................................................F..................................................V...................A.... 23ratNor ....................E..................................................................................F..................................................V...................A.... 24speTri ............H.......E................................................................................................................................F....V...................A.... 25dipOrd ..F....GT.GQG.....PEE...T........I........................................................................................................................V........................ 26cavPor .......P..N.H..R..Q.E...V.......................V...........................I..................R.............................V.................................V..............A.... 27oryCun ............H......E..............................................................................................................................R.......L...................A.... 28ochPri ....D.......H....F.........................................................................................L....D.....................------......R.......V...................A.... 29canFam .......R...........E........................................................I.............................................................................V...................A.... 30felCat .......P...........E.................................................--------------.......................................................................V...................A.... 31bosTau .......P.P...R.P........................................................V...I................................................................I............V.....A.............A.... 32turTru ......................K..........I.............................................................................V...................A.... 33susScr .......P.P...R.....E....................................................V...I.............................................................................V.....A.............A.... 34vicVic .......P.P...R.RH...............................L..................................................M........................................-------------------------------------.. 35equCab ............................................................................I.............................................................................V........................ 36myoLuc ............G.....Q.............................V.......................V...........................................................................T.....F........................ 37pteVam ......V.....H....V..............................V...........................I.............................................................................V........................ 38sorAra ............N..........................................................................................M.................................................VV...................A.... 39eriEur .S..Q.......G.........................................................................................................................................L...V...................A.... 40loxAfr .T.....P...D...Q..Q.....T.................................C..............................V................................................................V...................A.... 41proCap .T....V..E.D..S.........T................V......C.........Y..............................I...S..........................................H...-------------------------------------.. 42echTel .......P...NS...........V.........G.....I.................Y....................................S.......T..................................................V........................ 43dasNov ...........D...............................................K..............................................................................................V........................ 44choHof ......G....DS....F......................I....................R............................................................................................V.............L.......... 45monDom .....SVS...DYI..............................V...L.....I....K............V......................S.......V................................H....T....H..F....L.....T..A.V.FA.V.S.A.... 46macEug .V...L.....I....K............V..............................V................................H....T....H......VI.....T..A....A...N.A.... 47ornAna MT.YS.PQLGDY......E....V............V..V...V...L.....I.................V..................V...........M................................H....T....H.......I........A........N.A.... 48galGal ..SDCNSSS.E.Y....MQQE........R...II......V......L.....IF...K............V................S...F.S...I...M................................H.A..T....H..F....L.....T..A.V.FA.V.S.A.... 49taeGut ..SEYNNSS.E.YI....QEE........R...II.............L.....IF...K............V................S...F.S...M......C.............................H....T....H..F....I.....M..A.V.FA.V.S.A.... 50anoCar .EQGQNISS..DN----QQEE........V...I...V..LV......L.....I...TQ.....K......V................S..AF.S...I...S.............I..............F...H....T....H.VF...GI..S..A..A.I.FA.F.N.A.... 51xenTro ..G.SSYREESGYI...E..S........R...IF..V..MA......L.....I..ACS........................T....A.V...S.......NA..................L.V..........H.R..T....R..F.A..V.....TL.A.L....V.N.A.... 52danRer .E-NET-SISSGYI....LR.........K...I..A..ILV.....AT.....M..TFK..T..K.P....L...IF.F....S....F.V.S.S...L...Q...Y.................I..F.......H.R..T....H..FLSVVF.....A..A...V..W.N.A.... 53pimPro .E-NDT-SIPSGYV....LR.........K...I..A..ILV..V..AT.....I.QTIK..T..K.P.F..L....F.F....T....F.V.S.S...L...Q...Y.................I..........H.R..T.F..H..FL..VFT.L..A..A...V..W.N.A.... 54takRub .E-NET-WTHSSYV....LR......R..K...I..AL.IC...LM.AT........TFK..T..K.P.L..L...IF.F....T....F.V.SLS...L...E...F.................V..........H.R..A....H..FL...SV....A..A......W.S.A.... 54tetNig .D-NET-RSHPSYV....LR......R..K...I..A..IF...VM.AT........TFK..T..K.P.L..V...IF.F....T....F.V.SLS...L...E...F.................V..........H.R..T....H..FV...LV....A..A......W.S.A.... 56gasAcu .E-NET-WTHPSYI....LR......R..K...II.A..IC....M.AT.....I..TIK..S..K.P.L..V....F.F....T....FVV.S.A...L...E...F.................V..........H.R..T....Q..FL..VFV.M..A..A......W.N.A.... 57oryLat .E-N.S-W.HSSYV....LR......R..K...I..A..IL....M.AT.....I..TIK..S..K.P.L..V....F.F....T....FVV.S.S...L...E...F.................V..........H.R..T....Q.IFL..VFV.I..A..A......W.S.A.... 59calMil .TFDNSTALYSGYWL.DSLH....V........IISAC..IVT.L...L.....I.L.ITQ.R..K.P..LIT...IS.F.M..G.Q..L.....S...I...V....H................V..........H.Q..S..Q.R.VFMS..F..F..A..A......W.N.A.... 10homSap MALNHTALPQDERLPHYLRDGDPFASKLSWEADLVAGFYLTIIGILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGISVVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSYGVWLKRKHAYICLAAIWAYASFWTTMPLVGLGDYVPE.. phyloSNP ..................AA......B......A..A..B.....B.BB.........B......C.A.........B.A....A..........B......................................A...B..A.......A.............A............... .. .. position 1.1.........1.........2.........2.........2.........2.........2.........2.........2.........2.........2.........2.........3........3..........3.........3.........3.........3.....3 position 7.8.........9.........0.........1.........2.........3.........4.........5.........6.........7.........8.........9.........0........1..........2.........3.........4.........5.....6 position 89012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456 excMemCy eeeeeeeeeeeeeeeeeeeeMMMMMMMMMMMMMMMMMMMMMccccccccccccccccccccccccccccccccccMMMMMMMMMMMMMMMMMMMMMeeeeeeeeeeeeeeeMMMMMMMMMMMMMMMMMMMMMcccccccccccccccccccccccccccccccccccccccccccccc* keyResid ...diS................................................................................................................K........................................... exonNumb 44444444444444444444444444444444444444444444444444444444444444444444444444455555555555555555555555555555555555555555555555555555555555555555555555555555555556666666666666666666666 10homSap PFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTKVAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKS-LEGFRLHTVT-TVRKSSAVLEIHEEV* 11panTro .................................................................................................................................................E.....-..........-...............* 12gorGor ...............................................................P.....................................K.................................................-....Q.....-...............* 13ponPyg .....................................................................................................S.................................................-..........-...............* 14nomLeu .......................................................................................................................................................-..........-S..............* 15rheMac .......................................................................................................................................................-..........-...............* 16papHam .......................................................................................................................................................-..........-...............* 17calJac .......................................................................................................................................................-..D.......-...............* 18tarSyr ......................................................G.........T......................................................................................-..D.......-...............* 19otoGar ..-...........-.L...............................................G................................................................................T.....-..D.......-............Q..* 20micMur .......................................................I.............................................................................----------------------H......-A...........Q..* 21tupBel ....................I..................................I..........................................................................................................-............Q..* 22musMus ................G.......S.................A...........................V...............................................................R.....A...RG.....-..D.......-............Q..* 23ratNor ................G.......S.................A...........................V................................N..............................R.........R......-..D.......-A...........Q..* 24speTri .......................-..............E..............E..............-------.......................................S....................................-..D.......-A.......V...Q..* 25dipOrd ................LA.................S...........................P.......................................................................................-.......................Q..* 26cavPor ................A...I...H...........M..................I.............................................................................SR.....NA.........-..D.......-.D...-......Q..* 27oryCun ........................................................................................................................................S..R.S.........-..D.......-............Q..* 28ochPri ................................................................G.......................................................................S..R.....Q.....-..D.......-............Q..* 29canFam ................L...I.........................................................................................................................R........-..D...N...-............Q..* 30felCat ....................I..................................................................................................................................-..D.......-............Q..* 31bosTau ....................I.................................................V................................................................................-..D.......-..........V.Q..* 32turTru ....................I..........................................P..........................................................V............................-..D.......-............Q..* 33susScr .....................................................................................V.................................................................-..D.......-...........RQ..* 34vicVic ---------------------------------------------------------------------------............................................................................-..D....A..-............Q..* 35equCab ....................I...........................................G.....V................................................................................-..D.......-............Q..* 36myoLuc ...............T....I.................................K.........-----------.............................................S..............L........R......-..N.......-............Q..* 37pteVam ....................I...........................................G..M.........................................................................S..R......-..D.....I.-...EA.......Q..* 38sorAra ....................I..................................................................................N..............................R.....S...R......-.DD.......-...E........Q..* 39eriEur ................L...I.................................K............M..............................................................................N....-.KDY...................q..* 40loxAfr ....................I..................................I...........M............................................................................R......-..........-..K.......V.Q..* 41proCap ---------------------------------------------------------------------------..............................V.................................R.R..R...E..-...V......-............Q..* 42echTel .......................................................I...........M............................................................................................I.-...........HQ..* 43dasNov ................................................................................................................................................R......-..D.......-...E......V.Q..* 44choHof ...................................................................M.......................................................................R....R......-F.........-............Q..* 45monDom .................A..A.V.S.......F.............L.....T.....Y.T..QN..I.................................Q.....V.F.......................C......S..Q..A..E.-.RTY......-...R........Q..* 46macEug .................T..T...............................T..........Q..............................................................................................RHTVSTIRKSSSVSETYQ..* 47ornAna .................A..A...............................T..........QN....................................Q.......F.......................CRIS..RL..P.TG..E.-.KNS.S.SMS-.I..P...SGP.Q..* 48galGal .................A..A.V.S.......F.............L.....T.....Y.T..QN..I.................................Q...V...F.......................C.....RS..P.TLQ...S.KES.MY.IS-SH.D.A.LSGTQL..* 49taeGut .................A....V.S.....................L.....T.....Y.T..QN..I.................L...................V...F......................ECRL...RP..* 50anoCar ..............G..A..A..........V......M.C........Q..T.....Y.T..QNQ..................MF...................V..KV..I.............V......C.S...RP.N.QPLQ..NSR* 51xenTro ....T............K..I.V.SM......F..M......A.........A.......T.NQNN.T..I.................F............Q......E......MM....S...........C.P...KKD--.SLQNTT----S.VY.IS-.F...TTSAR* 52danRer ............T....S..S.VMCM.....IF..VI......M..F.....A...S...T.NKNN.S............................M....E...V..PV..........S............C.KK.VKSCCFQ.WR..KPSKTS.FY.ISGSIKQRPGD-.ASI.I* 53pimPro .................S..S.VMCM.I...V...G.......M..LQ....AQ..S...TQNKNK.H............................V....D...V..SI..........S............C.KN.AKLSCFQ.WS.RKHYKTS.FYSISASMK.RP.N-.VPT.I* 54takRub .................S..S.VMA......I...GI......M..F.....A..IS...A..RN..D..I.........................I....E...V..PV..I.......S..........V.V.TS.TNFSCC..L.ERIHFRKS..Y.ISGSL.DPLPPK.A.I.M* 54tetNig ....A............S..S.VMA......V...GI.....IM..F.....A..IS...A..KN..S..I.........................V....E...V..PV..I.......S..........A.L.TS.TSSSCC..L.ERVLFRKA..Y.ISGSL.DTLPPK.A.I.M* 56gasAcu .................S..S.VVA......V..AGI......M..F.....A..ISN..A..KN..N..I.........................V....E...V..SV..I.......S..........L.L.NS.MKSSCF.GL..PRHFRKS.FY.ISGS.KDNTTAK.AQI.M* 57oryLat .................S..S.VMS......V..AGI......M..C.........SS..A..KN..T..I.........................V....E......PV..I.......S.........LV.L.NS.-S.CCA.VIR.RTHFRNS.FY.ISGSLKDTAPAK.A.I.I* 59calMil .............RV..S.LI.V.T.........III.....I.........A..........QNH.S...N..........................................................................................................* 10homSap PFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTKVAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKS-LEGFRLHTVT-TVRKSSAVLEIHEEV* phyloSNP ...........................................B............B......CA.......................................................B.................................B...B.BBA............A..*
Neuropsin (NEUR1 compared to newropsin (NEUR2)
Newropsins are a new opsin gene family -- first reported here -- most closely related to neuropsins (42% percent identity) and next to melanopsins and peropsins. Like so many opsin families, they persist from chondrichthyes to archeosaurs but vanish without a trace in platypus, marsupials, and placentals. (The syntenic order B4GALT6 NEUR2 KIAA1012 remains conserved in mammals but no NEUR2 debris remains.) Newropsins retain many key attributes of GPCR signaling proteins and indeed opsins such as the seven transmembrane arrangement, Schiff base lysine, counterion tyrosine, amino terminal glycosylation site, and disulfide but have a very odd replacement of the G-protein binding site DRY with (invariantly conserved) VCC.
This motif must be an ancient derived feature that followed the gene duplication event with neuropsin since the much older DRY could not plausibly have re-evolved in neuropsin from VCC. Newropsins very likely link covalently via their orthologous Schiff base lysine with a retinal and interact with light according to some action spectrum. The VCC motif has been conserved over billions of years of branch length so cannot reflect simple loss of functionality; however its signaling capabilities if any are unclear.
It seems feasible that non-signaling opsins have become photoisomerases. Their substrate, while evidently involving an aldehyde capable of forming a Schiff base with the conserved lysine and likely interconverting cis/trans double bonds, need not be part of a ciliary opsin replenishment cycle. Other metabolic derivatives of beta-carotenes and lincopenes such as retinoic acid intermediates might be the substrates.
It must be recalled too that quite a diversity of photoreceptive retinoids are used in non-mammalian species, for example 9-cis isorhodopsin, porphyropsin (or 3-dehydroretinal vitamin A2) in freshwater fishes and some frogs, 3-dehydroretinal in freshwater crayfish, all-trans-5,6-dihydroretinal in cottoid fish in Lake Baikal and so forth. These are spectrally influenced by the surrounding opsin. Here too the negatively charged counterion, Glu113 in bovine rhodopsin, an alternative glutamate in melanopsins, or a special bound chloride ion. The counterparts of these are not known in neuropsins.
Below, conserved residues are shown for NEUR2 relative to human neuropsin (which represents that family accurately). Newropsin orthologs are rather rapidly diverging, especially in teleost fish. Transmembrane domains on newropsins were assigned by homology to neuropsin (taken from SwissProt ab initio annotation consistent with experimentally determined bovine rod rhodopsin). Newropsin is relatively truncated amino terminally but very extended in a highly variable manner carboxy terminally. That extension would lie in the cytoplasm and possibly be removed endoproteolytically. The early glycosylation is present in all species but appears shifted distally by four residues in tetrapods relative to fish.
According to transcript annotations, newropsin is expressed in zebrafish anterior segment (minus lens), fish brain and testes (Pimephales and Oncorhynchus), embryo, oviduct and fat body (Xenopus). These, while familiar sites from other opsins, provide only meagre constraints on possible newropsin functionality and association with photoreceptive tissues.
The intronation pattern is not a perfect match to neuropsin as might be expected. Some of the difference is lineage-specific, such as a gain in zebrafish, but other differences may be much older. Unless homologs have been retained recognizably in earlier diverging species, it won't be feasible to date the original gene duplication. No NEUR2 could be located in lamprey, tunicate, or lancelet.
Newropsin is further evidence that the neuropsin/peropsin/rgropsin group played a much greater role in ancestral vertebrate photoreception (which persisted into contemporary species), roles which were lost in stem mammals. That is quite similar to the ciliary opsin story. Overall mammals have retained less than half (7 of 17) of the vertebrate opsin repertoire. Such widespread gene loss is fully consistent with an old inference of a nocturnal era during which no selective pressure existed to maintain these photoreceptors.
position ...................................................................................................1.........1.........1.........1.........1.........1.........1.........1........1. position .........1.........2.........3.........4.........5.........6.........7.........8.........9.........0.........1.........2.........3.........4.........5.........6.........7........8. position 123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890 excMemCy eeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeMMMMMMMMMMMMMMMMMMMMMccccccccccccccccccccMMMMMMMMMMMMMMMMMMMMMeeeeeeeeeeeeeMMMMMMMMMMMMMMMMMMMMMcccccccccccccccccccccMMMMMMMMMMMMMMMMMMMMMeeeeeeee keyResid ....GLC.........glc.glc.................................................................................diS..cIon.................DRY?.............................................. NEUR2_galG MDPSFANST-FQSKITEAADIVVGTCYMVFGICSLCGNSILLYISYKKKHLLKPAEYFIINLAISDLAMTLTLYPLAVTSSLSHRWLYGKHICLFYAFCGLFFGICSLSTLTLLSVVCCLKICFPAYGNRFRRKHGQILIACAWTYAAIFACSPLAHWGEYGEEPY NEUR2_anoC MESYFANTT-FHSKITEAADVIVGVFYIVFGICSFCGNSILLYVSYKKKNLLKPAEYFMINLAISDLGMTLTLYPLAVTSSLAHRWLFGQQVCLFYAFCGVFFGVCSLTTLTLLSIVCCLKICFPVYGNRFRPGHGWILIACAWVYAAIFAFSPLAHWGEYGAEPY NEUR2_xenT MGNKSDASA-FYSSISETDDIVLGVLYSVFGLLSLSGNSMLLLVAYRKRSILKPAEFFIVNLSISDLGMTGTLFPLAIPSLFAHRWLFDKVTCNYYAFCGMLFGLCSLTNLTVLSSVCCLKVCYPAYGNKFSTAHSRILLLGIWAYAGLFATAPLADWGKYGPEPY NEUR2_danR MGNVSKTAL-FMSTISRQHDILMGSLYSVFFVLSLLGNGMLLFVAYRKRSSLKPAEFFVVNLSVSDLGMTLSLFPLAIPSALAHRWLFGEITCLCYAVCGVLFGLCSLTNLTALSSVCCLKVCFPNYGNKFSSSHACVMVIGVWCYASVFAVGPLVHWGSFGPEPY NEUR2_pimP MGNVSETAL-FVSTISRQHDILMGSLYSVFCVLSLLGNGMLLFVAYRKRSSLKPAEFFVINLSVSDLGMTLSLFPLAIPSALAHRWLFGEVVCLCYAVCGVLFGLCSLTNLTALSSVCCLKVCCPNYGNKFSSNHACVMVIGVWCYASVFAVGPLIRWGSFAPEPY NEUR2_tetN MGNASDTSDAFNSKISKEHDFLIGSIYSVFCVLSLMGNCILLLVAHHKRSTLKPAEFFIVNLSISDLGMTLTLFPLAIPSSFSHRWLFGEIACQLYATCGVLFGLCSLTNLTVLSSVCCLKVCLPNLGSKFSSSHARLLVAGVWGYASVFAVGPLVQWGHYGPEPY NEUR2_takR MGNASEASDIFLSKISKEHDILIGSIYSVFGLLSLAGNCILLLVAYHKRSMLKPAEFFIINLSISDLGMTLTLFPLAIPSSFSHRWLFGEITCQLYAMCGVLFGLCSLTNLTALSLVCCLKVCFPNHGSRFSSSHARLLVVGVWCYASVFAVGPLVQWGHYGPEPY NEUR2_gasA MGNASDTSAVFASTISKERDILMGSLYSVFGVLSLVGNCILLLVAYHKRSTLKPAEFFIINLSISDLGMTLSLFPLAIPSAFKHRWLFGELTCQLYAMCGVLFGLCSLTNLTALSFVCCLKVCFPNHGNRFSSSHARLLVVAVWGYASVFAVGPLARWGRYSPEPY NEUR2_oryL MGNVSDTSSLFASSISREHDILMGSLYSVFGLLSLSGNSMLLLVAYRKRSILKPAEFFIVNLSISDLGMTGTLFPLAIPSLFAHRWLFGEITCQLYAMCGVLFGLSSLTNLTALSLVCCLKVCFPNHGNKFSFSHARLLVAGVWCYASVFAVGPLARWGRYSAEPY NEUR2_calM GILSLVGNSVLLFVAYRKRQILKPAEYFVANLAVSDISMTVTLLPLAISSNFSHRWLFVSKPCMYYGFCSMLFGICSLTNLTVLSTVCCMKVCFPAYMSVVMIV-MFLLAWSPYSIVCLWASFGNPKLIPPAMAII NEUR1_homSa MALNHTALPQDERLPHYLRDGDPFASKLSWEADLVAGFYLTIIGILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGISVVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSYGVWLKRKHAYICLAAIWAYASFWTTMPLVGLGDYVPEPF NEUR1_canFa MALNHTARPQDERLPHYLREGDPFASKLSWEADLVAGFYLTIIGILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAICDLGISVVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSYGVWLKRKHAYICLAVIWAYASFWTTMPLVGLGDYAPEPF NEUR1_musMu MALNHTALPQDERLPHYLRDEDPFASKLSWEADLVAGFYLTIIGILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGISVVGKPFTIISCFCHRWVFGWFGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSYGVWLKRKHAYICLAVIWAYASFWTTMPLVGLGDYAPEPF NEUR1_loxAf MTLNHTAPPQDDRLPQYLQDGDPFTSKLSWEADLVAGFYLTIIGILSTFGNGYVLYMSCRRKKKLRPAEIMTINLAVCDLGISVVGKPFVIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSYGVWLKRKHAYICLAVIWAYASFWTTMPLVGLGDYAPEPF NEUR1_monDo MALNHSVSPQDDYIPHYLRDGDPFASKLSWEADLVAGFYLTIIGVLSTLGNGYVIYMSSKRKKKLRPAEIMTVNLAVCDLGISVVGKPFTIISCFSHRWVFGWVGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICHLSYGTWLKRHHAFICLALIWAYATFWATVPFAGVGSYAPEPF NEUR1_ornAn MTNYSAPQLGDYLPHYLREGDPFVSKLSWEADLVAGVYLVIIGVLSTLGNGYVIYMSSRRKKKLRPAEIMTVNLAVCDLGISVVGKPFTIVSCFCHRWVFGWMGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICHLSYGTWLKRHHAYICLAIIWAYASFWATMPLVGLGNYAPEPF NEUR1_calMi MTAFDNSTALYSGYWLHDSLHGDPFVSKLSWEADIISACYLIVTGLLSTLGNGYVIYLSITQKRKLKPPEILITNLAISDFGMSVGGQPFLIISCFSHRWIFGWVGCRWHGWAGFFFGCGSLITMTVVSLDRYLKICHLQYGSWLQRRHVFMSLAFIWFYAAFWATMPLVGWGNYAPEPF NEUR1_galGa MASDCNSSSQEEYLPHYMQQEDPFASKLSREADIIAGFYLTVIGILSTLGNGYVIFMSSKRKKKLRPAEIMTVNLAVCDLGISVVGKPFSIISFFSHRWIFGWMGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICHLAYGTWLKRHHAFICLALIWAYATFWATVPFAGVGSYAPEPF NEUR1_xenTr MAGNSSYREESGYIPHYERDSDPFASKLSREADIFAGVYLMAIGILSTLGNGYVIYMACSRKKKLRPAEIMTINLAVCDLGISVTGKPFAIVSCFSHRWVFGWNACRWYGWAGFFFGCGSLITLTVVSLDRYLKICHLRYGTWLKRRHAFIALAVIWAYATLWATLPLVGVGNYAPEPF NEUR1_danRe MENETSISSGYIPHYLLRGDPFASKLSKEADIVAAFYILVIGILSATGNGYVMYMTFKRKTKLKPPEIMTLNLAIFDFGISVSGKPFFIVSSFSHRWLFGWQGCRYYGWAGFFFGCGSLITMTIVSFDRYLKICHLRYGTWLKRHHAFLSVVFIWAYAAFWATMPVVGWGNYAPEPF NEUR1_takRu MENDTSIPSGYVPHYLLRGDPFASKLSKEADIVAAFYILVIGVLSATGNGYVIYQTIKRKTKLKPPEFMTLNLAVFDFGISVTGKPFFIVSSFSHRWLFGWQGCRYYGWAGFFFGCGSLITMTIVSLDRYLKICHLRYGTWFKRHHAFLCLVFTWLYAAFWATMPVVGWGNYAPEPF NEUR1_tetNi MENETWTHSSYVPHYLLRGDPFASRLSKEADIVAALYICIIGLMSATGNGYVLYMTFKRKTKLKPPELMTLNLAIFDFGISVTGKPFFIVSSLSHRWLFGWEGCRFYGWAGFFFGCGSLITMTVVSLDRYLKICHLRYGAWLKRHHAFLCLASVWAYAAFWATMPLVGWGSYAPEPF NEUR1_gasAc MDNETRSHPSYVPHYLLRGDPFASRLSKEADIVAAFYIFIIGVMSATGNGYVLYMTFKRKTKLKPPELMTVNLAIFDFGISVTGKPFFIVSSLSHRWLFGWEGCRFYGWAGFFFGCGSLITMTVVSLDRYLKICHLRYGTWLKRHHAFVCLALVWAYAAFWATMPLVGWGSYAPEPF NEUR1_oryLa MENETWTHPSYIPHYLLRGDPFASRLSKEADIIAAFYICIIGIMSATGNGYVIYMTIKRKSKLKPPELMTVNLAVFDFGISVTGKPFFVVSSFAHRWLFGWEGCRFYGWAGFFFGCGSLITMTVVSLDRYLKICHLRYGTWLKRQHAFLCLVFVWMYAAFWATMPLVGWGNYAPEPF NEUR1_pimPr MENTSWPHSSYVPHYLLRGDPFASRLSKEADIVAAFYILIIGIMSATGNGYVIYMTIKRKSKLKPPELMTVNLAVFDFGISVTGKPFFVVSSFSHRWLFGWEGCRFYGWAGFFFGCGSLITMTVVSLDRYLKICHLRYGTWLKRQHIFLCLVFVWIYAAFWATMPLVGWGSYAPEPF NEUR1_anoCa MEQGQNISSQDDNQQEEDPFASKLSVEADIVAGVYLLVIGILSTLGNGYVIYMSTQRKKKLKPAEIMTVNLAVCDLGISVVGKPFSIIAFFSHRWIFGWSGCRWYGWAGFFFGIGSLITMTAVSLDRYFKICHLSYGTWLKRHHVFICLGIIWSYAAFWATIPFAGFGNYAPEPF position 1.........1.........2.........2.........2.........2.........2.........2.........2.........2.........2.........2.........3........3..........3.........3.........3.........3.....3 position 8.........9.........0.........1.........2.........3.........4.........5.........6.........7.........8.........9.........0........1..........2.........3.........4.........5.....6 position 012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456 excMemCy eeeeeeeeeeeeeeeeeeMMMMMMMMMMMMMMMMMMMMMccccccccccccccccccccccccccccccccccMMMMMMMMMMMMMMMMMMMMMeeeeeeeeeeeeeeeMMMMMMMMMMMMMMMMMMMMMcccccccccccccccccccccccccccccccccccccccccccccc* keyResid .diS................................................................................................................K............................................................ NEUR2_galG GTACCIDWQSTNVDVMSMSYTVVLFVLCFILPCGVIVTSYSLILVTVKESRKAVEQHVSGPTRINNVQTITAKLSIAVCIGFFAAWSPYAIIAMWAAFGSIDKIPPLAFAIPAVFAKSSTLYNPIIHLLLKPNFRSNIAKDFTVIQQLCVR---CCFCVKELQ--TYRSTFNTGLRTFKG NEUR2_anoC GTACCIDWRISNMKKTAMSYTTALFVFCYIIPCGIIITSYTLILITVKDSRKAVEQHALGPTRMSSVHTITAKLSIAVCIGFFVAWSPYAIIAMWAAFGSIDMIPPLAFAVPAVFAKSSTLYNPAMYLFLKPNFRSTIAKDLTVLHRLCLK---SCFCPRGMQNCSYRSALEAPLKSFKG NEUR2_xenT GTACCLDWEASYRERKALSYTISLFVFCYLIPSSLIFISYTLIFVTVKGARRAVQQHLSPQAKGSSIHSLIIKLSIAVCIGFLIAWTPYAIVAMMAAFGDPTKIPSLVFALAAAFAKSSTIYNPVVYLLLKPNFLNVVTKDLTLFQTMCAV---VCGWCR-----TPAVKTPCPHKDLKT NEUR2_danR GTACCINWYTPSHDALAMSYIISLFIFCYVVPCTIIILSYTFILVTVRGSQQAVQQHVSPQTKVTNAHALIVKLSVAVCIGFLTAWSPYAIVAMWAAFSANEQVPPTAFALAAIMAKSSTIYNPMVYLLFKPNFRKSLSQDTQMFRHRICLSHSKASPSPGMKDQERQSSQQCNNKDGSI NEUR2_pimP GTACCINWYIPSHDALAMSYIISLFIFCYVVPCTIIILSYTFILLRVRGSRQAVQKHVSPKTKETNAHTLIVKLSVAVCIGFVTAWSPYAVVAMWAAFSANEPVPPTAFALAAILAKSSTIYNPMVYLLFKPNFRKILSQDTQNIRHRMCVSHSKASPTPEIK---AQSSQQC--KDATI NEUR2_tetN GTACCINWQAPNHELSSLSYIVCLFLFCYVLPCAIIILSYTCILMTVRGSRQAIQQHVSPQTKTANAHALIVKLSVAVCIGFLGAWSPYAVVAMWASFGDATWVPPDAFAIAAILAKSSTIYNPLVYLLCKPNFRECLYKDTSTLRQRIY----RGSPLSGPRDRSGGVTQR--HKDLSV NEUR2_takR GTACCIDWRAPNHELSSLSYIVCLFFFCYVLPCATIILSYTCILMTVRGSRQAIQQHVSPQTKTANAHSLIVKLSVAVCIGFLGAWSPYAIVAMWAAFGDATWVPPDAFAIAAILAKSSTIYNPVVYLLCKPNFRECLYKDTSTLRQRIY----RGSPQSEPRERFGGTSQR--HKDLSI NEUR2_gasA GTACCIDWHAPNHELAALSYIVCLFVFCYALPCATIFLSYTFILLTVRGSRQAVQQHVSPQTKTTNTHALIVKLSVAVCIGFLGAWTPYAVVAIWAAFGDATLVPPDAFALAAMFAKSSTIYNPVVYLLCKPNFRACLYRDTTLLRQRIY----RGSPRSEPKAHFGSTSQR--NKDMSV NEUR2_calM APLFAKSSTFYNPCIYVISYTMTVIAVNFVVPLSVMFFCYYNV NEUR2_oryL GTACCIDWHAPNHELWALSYILCLFIFCYALPCTIIFLSYAFILLTVRGSRQAVQQHVSPQTKTTNAHTLIVKLSVAVCIGFLGAWTPYAVIAMWAAFGDATQVPPTAFALAAVFAKSSTIYNPMVYLLCKPNFRECLCRDTSLLRHMIY----RGSP--QPQERFGSDSRR--NKDITA NEUR1_homSa GTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTKVAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKA-TKKKSLEGFRLHTVT-TVRKSSAVLEIHEEV NEUR1_calJa GTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTKVAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKA-TKKKSLEDFRLHTVT-TVRKSSAVLEIHEEV NEUR1_canFa GTSCTLDWWLAQASLGGQIFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTKVAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGRLKA-TKKKSLEDFRLNTVT-TVRKSSAVLEIHQEV NEUR1_musMu GTSCTLDWWLAQASGGGQVFILSILFFCLLLPTAVIVFSYAKIIAKVKSSSKEVAHFDSRIHSSHVLEVKLTKVAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYRFACCQAGGLRG-TKKKSLEDFRLHTVT-TVRKSSAVLEIHQEV NEUR1_loxAf GTSCTLDWWLAQASVGGQIFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEIAHFDSRIHSSHMLEMKLTKVAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLRA-TKKKSLEGFRLHTVT-TVKKSSAVLEVHQEV NEUR1_monDo GTSCTLDWWLAQASVAGQAFVLSILFFCLLFPTAVIVFSYVKIILKVKSSTKEVAHYDTRIQNSHILEMKLTKVAMLICAGFLIAWIPYAVVSVWSAFGQPDSIPVQFSVVPTLLAKSAAMYNPIIYQVIDCKFACCQSGGQKA-AKKESLRTYRLHTVT-TVRRSSAVLEIHQEV NEUR1_ornAn GTSCTLDWWLAQASVAGQAFILNILFFCLLLPTAVIVFSYVKIIAKVKSSTKEVAHFDSRIQNSHVLEMKLTKVAMLICAGFLIAWIPYAVVSVWSAFGQPDSIPIQFSVVPTLLAKSAAMYNPIIYQVIDCRISCCRLGGPKT-GKKESLKNSRSHSMS-TIRKPSAVSGPHQEV NEUR1_calMi GTSCTLDWWLARVSVSGLIFVLTILFFCLLLPIIIIVFSYIKIIAKVKSSAKEVAHFDSRIQNHHSLEMNLTK NEUR1_galGa GTSCTLDWWLAQASVAGQAFVLSILFFCLLFPTAVIVFSYVKIILKVKSSTKEVAHYDTRIQNSHILEMKLTKVAMLICAGFLIAWIPYAVVSVWSAFGQPDSVPIQFSVVPTLLAKSAAMYNPIIYQVIDCKFACCRSGGPKTLQKKSSLKESRMYTIS-SHRDSAALSGTQLEV NEUR1_xenTr GTTCTLDWWLAQASVKGQIFVLSMLFFCLLFPTMVIVFSYAKIIAKVKSSAKEVAHFDTRNQNNHTLEIKLTKVAMLICAGFLIAWFPYAVVSVWSAFGQPDSIPIELSVVPTMMAKSASMYNPIIYQVIDCKPACCKK------DKSLQNTTSRVYTIS-TFRKSTTSAR NEUR1_danRe GTSCTLDWWLTQASVSGQSFVMCMLFFCLIFPTVIIVFSYVMIIFKVKSSAKEVSHFDTRNKNNHSLEMKLTKVAMLICAGFLIAWIPYAVVSVMSAFGEPDSVPIPVSVVPTLLAKSSAMYNPIIYQVIDCKKKCVKSCCFQAWRKKKPSKTSRFYTISGSIKQR-PGDEASIEI NEUR1_takRu GTSCTLDWWLAQASVSGQSFVMCMLIFCLVLPTGVIVFSYVMIILQVKSSAQEVSHFDTQNKNKHHLEMKLTKVAMLICAGFLIAWIPYAVVSVVSAFGDPDSVPISISVVPTLLAKSSAMYNPIIYQVIDCKKNCAKLSCFQAWSKRKHYKTSRFYSISASMKKR-PANEVPTEI NEUR1_tetNi GTSCTLDWWLAQASVSGQSFVMAILFFCLILPTGIIVFSYVMIIFKVKSSAKEISHFDARIRNSHDLEIKLTKVAMLICAGFLIAWIPYAVVSVISAFGEPDSVPIPVSVIPTLLAKSSAMYNPIIYQVVDVKTSCTNFSCCKALKERIHFRKSRLYTISGSLRDPLPPKEAHIEM NEUR1_gasAc GTACTLDWWLAQASVSGQSFVMAILFFCLVLPTGIIVFSYIMIIFKVKSSAKEISHFDARIKNSHSLEIKLTKVAMLICAGFLIAWIPYAVVSVVSAFGEPDSVPIPVSVIPTLLAKSSAMYNPIIYQVADLKTSCTSSSCCKALKERVLFRKARLYTISGSLRDTLPPKEAHIEM NEUR1_oryLa GTSCTLDWWLAQASVSGQSFVVAILFFCLVLPAGIIVFSYVMIIFKVKSSAKEISNFDARIKNSHNLEIKLTKVAMLICAGFLIAWIPYAVVSVVSAFGEPDSVPISVSVIPTLLAKSSAMYNPIIYQVLDLKNSCMKSSCFKGLKKPRHFRKSRFYTISGSVKDNTTAKEAQIEM NEUR1_pimPr GTSCTLDWWLAQASVSGQSFVMSILFFCLVLPAGIIVFSYVMIICKVKSSSKEVSSFDARIKNSHTLEIKLTKVAMLICAGFLIAWIPYAVVSVVSAFGEPDSIPIPVSVIPTLLAKSSAMYNPIIYQLVDLKNSC-STCCAKVIRKRTHFRNSRFYTISGSLKDTAPAKEAHIEI NEUR1_anoCa GTSCTLDWWLAQGSVAGQAFILNILFFCLVLPTAVIMFCYVKIIAKVQSSTKEVAHYDTRIQNQHVLEMKLTKVAMLICAGFMFAWIPYAVVSVWSAFGRPDSVPIKVSVIPTLLAKSAAMYNPVIYQVIDCKSACCRPGNLQPLQKKNSR
Neuropsin (NEUR1 compared to newtopsin (NEUR3)
A third paralogous family NEUR3 was reported in July 2008 and characterized by syntenic relations and expression in chicken. However in chicken, NEUR1 actually is most abundant of the three paralogs in developing and early post-hatch neural retina, notably in differentiating ganglion cells and amacrine cells.
Recoverable sequences range from lamprey to shark to fish (where a further lineage-specific tandem duplication has occurred) to frog to sauropsids, with the gene again lost in all mammals including platypus. This locus exhibits an ancestral fusion of exon 2-3 relative to neuropsin and newropsin. The tandem duplication in ray-finned fish could easily be mistaken for a whole genome duplication effect -- however there is no sign of such an event for any of the 3 paralogs in any of the 5 fish with assembled genomes.
The NEUR3 group could possibly signal through heteromeric G protein. The DRY motif is a bit unusual, consisting of V/I R F/Y whereas the YNPxIY x aliphatic is fairly conventional. The Schiff base K is preserved in both NEUR3a and NEUR3b sequences. The counterion glutamate or chloride ion has not been determined.
The other oddity is NEUR1 and NEUR3 are on the same chromosome for all species examinable. They are separated by a million or so bp as well as other coding genes. This possibly represents an old tandem duplication that experienced subsequent rearrangements. On the whole, synteny has been quite well preserved for all NEUR genes:
NEUR3_galGal +CRISP +RHAG ..... +MUT -NEUR3 ..... +CDC5L -SUPT3H +RUNX2
NEUR3_anoCar -CRISP2 +RHAG ..... +Mut -NEUR3 ..... +CDC5L -SUPT3H +RUNX2
NEUR3_xenTro -CRISP2 -RHAG -PPHLN +MUT -NEUR3 ..... +CDC5L ..... .....
NEUR3a_danRer +XRN2 -TSTA3 +MGST3 +MUT -NEUR3a -NEUR3b -NAPB +DNMT3A .....
NEUR3a_tetNig ..... ..... ..... +MUT -NEUR3a -NEUR3b ..... ..... +RUNX2
NEUR1_galGal ..... +TNFRSF2 +CD2AP +GPR111 +NEUR1 ..... +MRPL19 ..... .....
NEUR1_anoCar -GPR111 -TNFRSF2 +CD2AP ..... +NEUR1 -SPATS1 +MRPL19 ..... +ITSN2
NEUR1_xenTro ..... +TNFRSF21 +CD2AP +GPR111 +NEUR1 -PTCHD1 +MRPL19 ..... +ITSN2
NEUR1_danRer +GPR111 -TNFRSF21 +CD2AP ..... +NEUR1 +CNIH2 -LBR -ENAH +ITSN2
NEUR2_galGal -DSC1 +DSG2 +TTR -B4GALT6 -NEUR2 -K1012 +RNF138 ..... .....
NEUR2_anoCar -DSC1 +DSG2 +TTR -B4GALT6 -NEUR2 -K1012 +RNF138 ..... .....
A proposed revised terminology for this family follows. Note NEUR2 and NEUR3 will never receive official HGCN nomenclature because (like thousands of amniote ancestral genes) they are absent in human and mouse. Here lower case a and b are used in the case of lineage-specific duplications, with a reserved for the copy with higher blastp score to human (or, if absent, nearest species).
Gene Protein HGCN Synonyms Lineage-specific duplicate DRY YNPxxY K Accessions
NEUR1 neuropsin OPN5 cOpn5m NEUR1a/b cephalochordate DRY YNPIIY K NM_181744 NM_001130743
NEUR2 newropsin ---- cOpn5L1 VCC YNPxIY K XM_419178
NEUR3 newtopsin ---- cOpn5L2 NEUR3a/b actinopterygii vRf YNPxIY K XM_420056
NEUR4: a fourth neuropsin from lamprey to platypus
Yet another new opsin in this group! These genes were first described here on 29 Jan 2009 (note GenBank had frog gene correctly predicted but chicken gene chimerized in a misassembly). Like many opsins, NEUR4 orthologs range throughout the vertebrates with the exception of theran mammals. Platypus is thus again distinguished by its retention of this ancient gene, whereas it is long gone from marsupials and placentals. This pattern of retention is consistent with platypus being more bird-like than mammal and supports the Wall 'dark era' apparently experienced by mammals during which Monotremata were somehow less dramatically affected.
This opsin is unusual for its TRY at the DRY motif, though its Schiff base lysine region is standard (KSASFYNPIIYFGMNSKFR). Its best match within opsins is to NEUR1; outside of neuropsins to peropsins, melanopsins and various ancestral ciliary opsins (it has no special affinities to RGR). Transcripts are abundant in frog and fish and the former demonstrate expression in adult eye (ES678087). This opsin is clearly capable of signaling through some heterotrimeric Galpha protein but its function is unknown.
NEUR4 shares some early intron positions and phases with NEUR1 but otherwise the pattern differs significantly, suggesting rather ancient divergence after origination by segmental duplication (not as processed pseudogene subsequently re-intronated). This is consistent with a very low percent identity (38%) to NEUR1, considering that a 'floor' of about 25% identity relates any pair of GPCR (eg NEUR4 is 23% identical to its best non-opsin match in human, tachykinin receptor).
The phylogenetic distribution of the neuropsin gene swarm is puzzling. Only NEUR1 is found in non-vertebrate deuterostomes today yet family origins must be far more ancient as it is basal to ciliary opsins already found in cnidaria. Clearly neuropsins persisted for a very long period in pre-bilateran and bilateran ancestors prior to being lost (perhaps in a few stem events). Neuropsins may be important only in lineages where ciliary imaging opsins are important, yet all but NEUR1 have been lost in placentals despite persistence of the other three classes for several hundred million years.
Alternatively, the neuropsin gene family expanded in the lamprey stem, diverged rapidly in primary sequence, and experienced multiple intron reorganization events. Because even one is rare, two independent events is rare-squared, and three is effectively impossible. Because the minimal number of events needed to synchronize intronation is five, this hypothesis appears unsustainable. Each orthology class has the same intronation pattern in all its members back to the earliest divergence for which sequence is available.
Introns compared in four neuropsins 1^2 etc indicate relative intron gain. Exons 1 is omitted (same in all 4 genes). Irrelevant amino acids not shown (...) NEUR1 GIL...GIS 1^2 VVG...SHR WIF...YGW AGF...LAY GTW...SYA PEP...LTK ESR 2^1 MYT...LEV* NEUR3 AIL...GMA ISM...NHA WLG...YAL MGF...KSN SNK...YYG PEP...LTL SAD NSA...ARH* NEUR2 GIC...AMT LTL...SHR 2^1 WLY...YAF CGL...PAY GNR...EYG EEP...TAK RCC FCV...TDL* NEUR4 GWM...SIS VFG...NVF RDD...DGF 0^0 LTL...PER AHC...SYT 1^2 DRM...VTR KLK RFK...DRL*
The most parsimonious explanation here -- assuming the gene tree (((NEUR1, NEUR3), NEUR2), NEUR4) -- is that NEUR3 represents the ancestral intronation pattern. This correlates well with its relative lack of retroposon events which may facilitate intron gain -- a single unshared CR1 LINE -- and the gene span is 1/3 to 1/8 of the others. NEUR1 has 7 LINE elements for comparison. NEUR1, NEUR2 and NEUR4 separately acquired new introns (different locations and phases) within the second exon. NEUR1 also acquired a new intron in the terminal exon (which curiously is alignable to the end only in NEUR3). Intron gain is both rare in vertebrate coding genes and considerably less common than intron loss, yet the events here are more economically placed on terminal gene tree leaves as intron gains.
NEUR1 and NEUR3 both lie on the minus strand of chicken chromosome 3, separated by 1,310,977 bp and a half dozen coding genes. While not adjacent, this still suggests recent tandem duplication followed by local inversions, which is supported by the deeper match of the run-on terminal exon. This arrangement is readily tracked back to teleost fish, indicating the last expansion of neuropsin occurred prior to this, yet not by whole genome duplication. Perhaps the genes will be directly tandem in the upcoming revised Callorhinchus and Petromyzon genomes.
In chicken but not lizard, NEUR4 also lies on this same chromosome, though greatly distant. This possibly indicates that it too arose from tandem duplication of NEUR1 though at a much earlier date. More likely its current position is coincidental because, while chickens have 39 chromosomes, only 6 are macro-sized.
Indels are another type of potentially informative rare genomic event. Upon alignment of the reference sequence set below, five phylogenetically coherent indels emerge. Amphioxus and sea urchin genes can be used as outgroup to determine ancestral length (eg, to resolve each indel as deletion or insertion); this gives the same outcome as alignment to all ciliary opsins and indeed many GPCR. Each indel affects every member of its orthology class and none affects more than one class (other than in fish where the event affected the parent NEUR3 gene prior to its tandem duplication). Thus, as with introns, indels are predominantly ancient and do not provide internal clustering of neuropsins genes to guide the gene tree. Possibly the pre-lamprey indels were fixed shortly after gene duplication as the new paralogs re-functionalized.
Indel #res type affected_genes timing location A 5 insert NEUR4 pre-lamprey 2 residues before first disulfide cysteine B 1 insert NEUR3 NEUR3a NEUR3b pre-teleost 2 residues after DRY motif, cytoplasmic loop C2 C 2 deletion NEUR4 pre-lamprey 1 residue before second disulfide cysteine D 2 deletion NEUR3b post-fish tandem 9 residues after second disulfide E 3 insert NEUR4 pre-lamprey 16 residues before Schiff lysine
It appears very likely NEUR1 is the parent gene providing the core (unknown) function:
- NEUR1 is best-blastp to amphioxus and sea urchin homologs whereas the others are undetectable outside vertebrates;
- NEUR1 survived the longest in mammals;
- NEUR1 is the best-Blastp for each of the others;
- NEUR1 alone retains the ancestral DRY signaling motif.
- NEUR1 has ancestral length (no indels relative to broader opsins or GPCR
Curated collection of neuropsins
These genes, almost all full length, have been extracted from various genome projects and cDNA data sets. In a few instances, accurate gene models had been previously computed by bioinformatic pipelines but these are so mixed with erroneous and mislabeled predictions as to be worthless for comparative genomics. The UCSC 44-species alignment is very helpful for rapid collection of individual exons taking care to note small insertions relative to human sequence are suppressed. The proteins below are parsed into exons whose coding phase is also shown. Because intronation is exceedingly conservative, genomically deduced introns can be reliable transfered to cDNA-only species.
Despite the incomplete nature of many genome assemblies and lack of transcripts in specialized cell types, the absence of a given neuropsin in a given clade is rarely attributable to lack of data. For example, while sloth assembly alone might have only 2x mean coverage (and thus lack many coding genes), overall coverage of its clade (Atlantogenata: armadillo, sloth, elephant, mammoth, hyrax, and tenrec) approaches 30x (90 million 1 kbp traces). Here coding genes can be missing only for the compositionally oddest or most extreme chromosomal locations.
For early diverging deuterostomes, the situation is somewhat different. Here only one species per divergence node is generally available and assemblies encountered extreme diploid heterozygosity and retroposon issues with little outside support from transcript programs (except for Ciona and sea urchin). The species chosen may be quite specialized within its clade and have experienced very extensive gene loss, much as Drosophila is wholly unrepresentative of the protostome genome. Lamprey, despite 19 million traces, has poor coverage with contigs rarely encompassing more than an exon or two.
However gene divergence (to the point of unrecognizability) is not an issue because the divergence floor to GPCR is well within tblastn reporting capabilities. Even in short contig assemblies, individual exons can be identified and reliably assigned to opsin orthology class using the classifier, provided the exon is reasonably conserved. Even poorly conserved N- and C-terminal exons can be extended outward to an initial methionine or stop codon with some reliability.
Consequently the phylogenetic 'end points' of a given gene are fairly certain even on the early-diverging side, though that remains somewhat muddled because lineage-specific loss is not at all uncommon in opsins or specialized species.
NEUR1: 52 deuterostome neuropsins
>NEUR1_homSap OPN5 1_6 43 0 1 2_6 40 2 1 3_6 57 2 1 4_6 112 2 0 5_6 81 0 2 6_6 19 1 0 0 MALNHTALPQDERLPHYLRDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAAIWAYASFWTTMPLVGLGDYVPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKSLEGFR 2 1 LHTVTTVRKSSAVLEIHEEV* 0 >NEUR1_panTro 0 MALNHTALPQDERLPHYLRDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAAIWAYASFWTTMPLVGLGDYVPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKETKKKSLEGFR 2 1 LHTVTTVRKSSAVLEIHEEV* 0 >NEUR1_gorGor 0 MALNHTALPQDERLPHYLRDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 12 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 YASFWTTMPLVGLGDYVPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIPSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGKPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKSLEGFQ 2 1 LHTVTTVRKSSAVLEIHEEv* 0 >NEUR1_ponPyg 0 MALNHTALPQDERLPHYLRDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAAIWAYASFWTTMPLVGLGDYVPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGSPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKSLEGFR 2 1 LHTVTTVRKSSAVLEIHEEV* 0 >NEUR1_nomLeu 0 MALNHTALPQDERLPHYLRDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAAIWAYASFWTTMPLVGLGDYVPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKSLEGFR 2 1 LHTVTSVRKSSAVLEIHEEv* 0 >NEUR1_macMul 0 MALNHTALPQDERLPHYLRDGDPFASKLSWEADIVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAAIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKSLEGFR 2 1 LHTVTTVRKSSAVLEIHEEV* 0 >NEUR1_papHam 0 MALNHTALPQDERLPHYLRDGDPFASKLSWEADIVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAAIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKSLEGFR 2 1 LHTVTTVRKSSAVLEIHEEv* 0 >NEUR1_calJac 0 MALNHTSLPQDERLPHYLRDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAAIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKSLEDFR 2 1 LHTVTTVRKSSAVLEIHEEV* 0 >NEUR1_tarSyr 0 MALNHTALPQDERLPHYLRDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VIGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAAIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKGVAHFDSRIHTSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKSLEDFR 2 1 LHTVTTVRKSSAVLEIHEEv* 0 >NEUR1_otoGar 0 MALNHTALPQDELRPHYLRDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTVNLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 WTMMPLVGLEDYVPEPFTSCTLDWWLAQSLGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHGSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKTTKKKSLEDFR 2 1 LHTVTTVRKSSAVLEIHQEV* 0 >NEUR1_micMur 0 MALNHTVLPQDERLPHYLRDGDPFASKLSWEADLVAGFYLIII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 1 2 GVWLKRKHAYICLALIWAYVSFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEIAHFDSRIHSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDHR 2 1 LHTVTAVRKSSAVLEIHQEv* 0 >NEUR1_tupBel 0 MALNHTALPQDESLPHYLRDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAVIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASVGGQIFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEIAHFDSRIHSSHVLEMKLTK 0 0 2 1 LHTVTTVRKSSAVLEIHQEV* 0 >NEUR1_musMus 0 MALNHTALPQDERLPHYLRDEDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWFGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAVIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASGGGQVFILSILFFCLLLPTAVIVFSYAKIIAKVKSSSKEVAHFDSRIHSSHVLEVKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYRFACCQAGGLRGTKKKSLEDFR 2 1 LHTVTTVRKSSAVLEIHQEV* 0 >NEUR1_ratNor 0 MALNHTALPQDERLPHYLRDEDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWFGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 gVWLKRKHAYICLAVIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASGGGQVFILSILFFCLLLPTAVIVFSYAKIIAKVKSSSKEVAHFDSRIHSSHVLEVKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPNSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYRFACCQTGGLRATKKKSLEDFR 2 1 LHTVTAVRKSSAVLEIHPEv* 0 >NEUR1_speTri 0 MALNHTALPQDEHLPHYLRDEDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAFICLAVIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASVGGQVFINILFFCLLLPTAVIEFSYVKIIAKVKSSSEEVAHFDSRIHSSHV 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPSLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKSLEDFR 2 1 LHTVTAVRKSSAVVEIHQEv* 0 >NEUR1_dipOrd 0 MAFNHTAGTQGQGLPHYLPEEDPFTSKLSWEADIVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAVIWAYASFWTTMPLVGLGDYVPEPFGTSCTLDWWLAQASLAGQVFILNILFFCLLLPTSVIVFSYVKIIAKVKSSSKEVAHFDSRIPSSHVLEMKLTK 0 0 2 1 * 0 >NEUR1_cavPor 0 MALNHTAPPQNEHLPRYLQDEDPFVSKLSWEADLVAGFYLTII 1 2 GILSTVGNGYVLYMSSRRKKKLRPAEIMTINLAICDLGIS 1 2 vVGKPFTIISCFRHRWVFGWIGCRWYGWAGFFFGCGSLITMTVVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAAIWAYVSFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASAGGQIFILHILFFCLLLPTAMIVFSYVKIIAKVKSSSKEIAHFDSRIHSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDSRFACCQNAGLKATKKKSLEDFR 2 1 LHTVTTDRKSAVLEIHQEV* 0 >NEUR1_oryCun 0 MALNHTALPQDEHLPHYLREGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRRHAYICLALIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFSCCRTSGLKATKKKSLEDFR 2 1 LHTVTTVRKSSAVLEIHQEv* 0 >NEUR1_ochPri 0 MALNDTALPQDEHLPHYFRDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRLYGWADFFFGCGSLITMTAVSLDRYLK 1 2 GVWLKRRHAYICLAVIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHGSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFSCCRTGGLKQTKKKSLEDFR 2 1 LHTVTTVRKSSAVLEIHQEv* 0 >NEUR1_canFam 0 MALNHTARPQDERLPHYLREGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAICDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAVIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASLGGQIFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGRLKATKKKSLEDFR 2 1 LNTVTTVRKSSAVLEIhQEV* 0 >NEUR1_felCat 0 MALNHTAPPQDERLPHYLREGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAE 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAVIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASVGGQIFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKSLEDFR 2 1 LHTVTTVRKSSAVLEIHQEv* 0 >NEUR1_bosTau 0 MALNHTAPPPDERRPPYLRDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTVNLAICDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GIWLKRKHAYICLAVIWAYAAFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASVGGQIFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEVKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKSLEDFR 2 1 LHTVTTVRKSSAVLEVHQEv* 0 >NEUR1_turTru 0 1 2 GILSTFGNGYVLYMSSRRKKKLKPAEIMTINLAICDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAVIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASVGGQIFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIPSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAVYNPIIYQVIDYKFACCQTGGLKATKKKSLEDFR 2 1 LHTVTTVRKSSAVLEIHQEv* 0 >NEUR1_susScr 0 MALNHTAPPPDERRPHYLREGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTVNLAICDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAVIWAYAAFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTK 0 0 VAMLICAGFLVAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKSLEDFR 2 1 LHTVTTVRKSSAVLEIRQEV* 0 >NEUR1_vicVic 0 MALNHTAPPPDERRPRHLRDGdPFASKLSWEADLVAGFYLTII 1 2 GILSTLGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWMFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKSLEDFR 2 1 LHAVTTVRKSSAVLEIHQEV* 0 >NEUR1_equCab 0 MALNHTALPQDERLPHYLRDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAICDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAVIWAYASFWTTMPLVGLGDYVPEPFGTSCTLDWWLAQASVGGQIFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHGSHVLEVKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKATKKKSLEDFR 2 1 LHTVTTVRKSSAVLEIHQEV* 0 >NEUR1_myoLuc 0 MALNHTALPQDEGLPHYLQDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTVGNGYVLYMSSRRKKKLRPAEIMTVNLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHTYICLAFIWAYASFWTTMPLVGLGDYVPEPFGTSCTLDWWLAQATVGGQIFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKKVAHFDSRIH 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSSAMYNPIIYQVIDYKLACCQTGGLRATKKKSLENFR 2 1 LHTVTTVRKSSAVLEIHQEv* 0 >NEUR1_pteVam 0 MALNHTVLPQDEHLPHYVRDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTVGNGYVLYMSSRRKKKLRPAEIMTINLAICDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAVIWAYASFWTTMPLVGLGDYVPEPFGTSCTLDWWLAQASVGGQIFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHGSHMLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTSGLRATKKKSLEDFR 2 1 LHTITTVREASAVLEIHQEV* 0 >NEUR1_sorAra 0 MALNHTALPQDENLPHYLRDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWMGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLVVIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASVGGQIFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPNSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYRFACCQSGGLRATKKKSLDDFr 2 1 LHTVTTVRESSAVLEIHQEV* 0 >NEUR1_eriEur 0 MSLNQTALPQDEGLPHYLRDGDPFASKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSSRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 gVWLKRKHAYLCLAVIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASLGGQIFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKKVAHFDSRIHSSHMLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLKANKKKSLKDYR 2 1 * 0 >NEUR1_loxAfr 0 MTLNHTAPPQDDRLPQYLQDGDPFTSKLSWEADLVAGFYLTII 1 2 GILSTFGNGYVLYMSCRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFVIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAVIWAYASFWTTMPLVGLGDYAPEPFGTSCTLDWWLAQASVGGQIFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEIAHFDSRIHSSHMLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLRATKKKSLEGFR 2 1 LHTVTTVKKSSAVLEVHQEv* 0 >NEUR1_proCap 0 MTLNHTVLPEDDRLSHYLRDGDPFTSKLSWEADLVAGFYLTVI 1 2 GILSTCGNGYVLYMSYRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFIIISSFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICHLSY 1 2 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSVPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCRTRGLRATKEKSLEGVR 2 1 LHTVTTVRKSSAVLEIHQEv* 0 >NEUR1_echTel 0 MALNHTAPPQDNSLPHYLRDGDPFVSKLSWEADLGAGFYLIII 1 2 GILSTFGNGYVLYMSYRRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFSHRWVFGWTGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAVIWAYASFWTTMPLVGLGDYVPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEIAHFDSRIHSSHMLEMKLTK 0 0 2 1 LHTITTVRKSSAVLEIHQEV* 0 >NEUR1_dasNov 0 MALNHTALPQDDRLPHYLRDGDPFASKLSWEADLVAGFYLTII 1 2 gILSTFGNGYVLYMSSKRKKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAVIWAYASFWTTMPLVGLGDYVPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCQTGGLRATKKKSLEDFR 2 1 LHTVTTVRESSAVLEVHQEV* 0 >NEUR1_choHof 0 MALNHTGLPQDDSLPHYFRDGDPFASKLSWEADLVAGFYLIII 1 2 GILSTFGNGYVLYMSSRRRKKLRPAEIMTINLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWIGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICYLSY 1 2 GVWLKRKHAYICLAVIWAYASFWTTMPLLGLGDYVPEPFGTSCTLDWWLAQASVGGQVFILNILFFCLLLPTAVIVFSYVKIIAKVKSSSKEVAHFDSRIHSSHMLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGRPDSIPIQLSVVPTLLAKSAAMYNPIIYQVIDYKFACCRTGGLRATKKKSFEGFR 2 1 LHTVTTVRKSSAVLEIHQEv* 0 >NEUR1_monDom 0 MALNHSVSPQDDYIPHYLRDGDPFASKLSWEADLVAGFYLTII 1 2 GVLSTLGNGYVIYMSSKRKKKLRPAEIMTVNLAVCDLGIS 1 2 VVGKPFTIISCFSHRWVFGWVGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICHLSY 1 2 GTWLKRHHAFICLALIWAYATFWATVPFAGVGSYAPEPFGTSCTLDWWLAQASVAGQAFVLSILFFCLLFPTAVIVFSYVKIILKVKSSTKEVAHYDTRIQNSHILEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGQPDSIPVQFSVVPTLLAKSAAMYNPIIYQVIDCKFACCQSGGQKAAKKESLRTYR 2 1 LHTVTTVRRSSAVLEIHQEv* 0 >NEUR1_macEug 0 1 2 GVLSTLGNGYVIYMSSKRKKKLRPAEIMTVNLAVCDLGIS 1 2 VVGKPFTIISCFCHRWVFGWVGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICHLSy 1 2 GTWLKRHHAYICLVIIWAYATFWATMPLAGLGNYAPEPFGTSCTLDWWLAQASVTGQTFILNILFFCLLLPTAVIVFSYVKIIAKVKSSTKEVAHFDSRIQSSHVLEMKLTK 0 0 2 1 RHTVSTIRKSSSVSETYQEV* 0 >NEUR1_ornAna 0 MTNYSAPQLGDYLPHYLREGDPFVSKLSWEADLVAGVYLVII 1 2 GVLSTLGNGYVIYMSSRRKKKLRPAEIMTVNLAVCDLGIS 1 2 VVGKPFTIVSCFCHRWVFGWMGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICHLSY 1 2 GTWLKRHHAYICLAIIWAYASFWATMPLVGLGNYAPEPFGTSCTLDWWLAQASVAGQAFILNILFFCLLLPTAVIVFSYVKIIAKVKSSTKEVAHFDSRIQNSHVLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGQPDSIPIQFSVVPTLLAKSAAMYNPIIYQVIDCRISCCRLGGPKTGKKESLKNSR 2 1 SHSMSTIRKPSAVSGPHQEV* 0 >NEUR1_galGal 0 MASDCNSSSQEEYLPHYMQQEDPFASKLSREADIIAGFYLTVI 1 2 GILSTLGNGYVIFMSSKRKKKLRPAEIMTVNLAVCDLGIS 1 2 VVGKPFSIISFFSHRWIFGWMGCRWYGWAGFFFGCGSLITMTAVSLDRYLKICHLAY 1 2 GTWLKRHHAFICLALIWAYATFWATVPFAGVGSYAPEPFGTSCTLDWWLAQASVAGQAFVLSILFFCLLFPTAVIVFSYVKIILKVKSSTKEVAHYDTRIQNSHILEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVWSAFGQPDSVPIQFSVVPTLLAKSAAMYNPIIYQVIDCKFACCRSGGPKTLQKKSSLKESR 2 1 MYTISSHRDSAALSGTQLEV* 0 >NEUR1_taeGut 0 MASEYNNSSQEEYIPHYLQEEDPFASKLSREADIIAGFYLTII 1 2 GILSTLGNGYVIFMSSKRKKKLRPAEIMTVNLAVCDLGIS 1 2 VVGKPFSIISFFSHRWMFGWIGCCWYGWAGFFFGCGSLITMTAVSLDRYLKICHLSY 1 2 GTWLKRHHAFICLAIIWAYAMFWATVPFAGVGSYAPEPFGTSCTLDWWLAQASVAGQVFVLSILFFCLLLPTAVIVFSYVKIILKVKSSTKEVAHYDTRIQNSHILEMKLTK 0 0 VAMLICAGFLLAWIPYAVVSVWSAFGRPDSVPIQFSVVPTLLAKSAAMYNPIIYQVIECRLACCRPGGCCRPGGLKAKSSLKKSR 2 1 TYTISAHRDSTAMNETQLEA* 0 >NEUR1_anoCar 0 MEQGQNISSQDDNQQEEDPFASKLSVEADIVAGVYLLVI 1 2 GILSTLGNGYVIYMSTQRKKKLKPAEIMTVNLAVCDLGIS 1 2 VVGKPFSIIAFFSHRWIFGWSGCRWYGWAGFFFGIGSLITMTAVSLDRYFKICHLSY 1 2 GTWLKRHHVFICLGIIWSYAAFWATIPFAGFGNYAPEPFGTSCTLDWWLAQGSVAGQAFILNILFFCLVLPTAVIMFCYVKIIAKVQSSTKEVAHYDTRIQNQHVLEMKLTK 0 0 VAMLICAGFMFAWIPYAVVSVWSAFGRPDSVPIKVSVIPTLLAKSAAMYNPVIYQVIDCKSACCRPGNLQPLQKKNSR 2 1 >NEUR1_xenTro 0 MAGNSSYREESGYIPHYERDSDPFASKLSREADIFAGVYLMAI 1 2 GILSTLGNGYVIYMACSRKKKLRPAEIMTINLAVCDLGIS 1 2 VTGKPFAIVSCFSHRWVFGWNACRWYGWAGFFFGCGSLITLTVVSLDRYLKICHLRY 1 2 GTWLKRRHAFIALAVIWAYATLWATLPLVGVGNYAPEPFGTTCTLDWWLAQASVKGQIFVLSMLFFCLLFPTMVIVFSYAKIIAKVKSSAKEVAHFDTRNQNNHTLEIKLTK 0 0 VAMLICAGFLIAWFPYAVVSVWSAFGQPDSIPIELSVVPTMMAKSASMYNPIIYQVIDCKPACCKKDKSLQNTTSR 2 1 VYTISTFRKSTTSAR* 0 >NEUR1_danRer 0 MENETSISSGYIPHYLLRGDPFASKLSKEADIVAAFYILVI 1 2 GILSATGNGYVMYMTFKRKTKLKPPEIMTLNLAIFDFGIS 1 2 VSGKPFFIVSSFSHRWLFGWQGCRYYGWAGFFFGCGSLITMTIVSFDRYLKICHLRY 1 2 gTWLKRHHAFLSVVFIWAYAAFWATMPVVGWGNYAPEPFGTSCTLDWWLTQASVSGQSFVMCMLFFCLIFPTVIIVFSYVMIIFKVKSSAKEVSHFDTRNKNNHSLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVMSAFGEPDSVPIPVSVVPTLLAKSSAMYNPIIYQVIDCKKKCVKSCCFQAWRKKKPSKTSR 2 1 FYTISGSIKQRPGDEASIEI* 0 >NEUR1_takRub 0 MENDTSIPSGYVPHYLLRGDPFASKLSKEADIVAAFYILVI 1 2 GVLSATGNGYVIYQTIKRKTKLKPPEFMTLNLAVFDFGIS 1 2 VTGKPFFIVSSFSHRWLFGWQGCRYYGWAGFFFGCGSLITMTIVSLDRYLKICHLRY 1 2 GTWFKRHHAFLCLVFTWLYAAFWATMPVVGWGNYAPEPFGTSCTLDWWLAQASVSGQSFVMCMLIFCLVLPTGVIVFSYVMIiLQVKSSAQEVSHFDTQNKNKHHLEMKLTK 0 0 VAMLICAGFLIAWIPYAVVSVVSAFGDPDSVPISISVVPTLLAAKSSAMYNPIIYQVVDVKTSCTNFSCCKALKERIHFRKSR 2 1 FYSISASMKKRPANEVPTEI* 0 >NEUR1_tetNig 0 MENETWTHSSYVPHYLLRGDPFASRLSKEADIVAALYICII 1 2 gLMSATGNGYVLYMTFKRKTKLKPPELMTLNLAIFDFGIS 1 2 VTGKPFFIVSSLSHRWLFGWEGCRFYGWAGFFFGCGSLITMTVVSLDRYLKICHLRY 1 2 GAWLKRHHAFLCLASVWAYAAFWATMPLVGWGSYAPEPFGTSCTLDWWLAQASVSGQSFVMAILFFCLILPTGIIVFSYVMIIFKVKSSAKEISHFDARIRNSHDLEIKLTK 0 0 VAMLICAGFLIAWIPYAVVSVISAFGEPDSVPIPVSVIPTLLAKSSAMYNPIIYQVADLKTSCTSSSCCKALKERVLFRKSR 2 1 YTISGSLRDTLPPKEAHIEM* 0 >NEUR1_gasAcu 0 MDNETRSHPSYVPHYLLRGDPFASRLSKEADIVAAFYIFII 1 2 GVMSATGNGYVLYMTFKRKTKLKPPELMTVNLAIFDFGIS 1 2 VTGKPFFIVSSLSHRWLFGWEGCRFYGWAGFFFGCGSLITMTVVSLDRYLKICHLRY 1 2 GTWLKRHHAFVCLALVWAYAAFWATMPLVGWGSYAPEPFGTACTLDWWLAQASVSGQSFVMAILFFCLVLPTGIIVFSYIMIIFKVKSSAKEISHFDARIKNSHSLEIKLTK 0 0 VAMLICAGFLIAWIPYAVVSVVSAFGEPDSVPIPVSVIPTLLAKSSAMYNPIIYQVLDLKNSCMKSSCFKGLKKPRHFRKSR 2 1 YTISGSLRDTLPPKEAHIEM* 0 >NEUR1_oryLat 0 MENETWTHPSYIPHYLLRGDPFASRLSKEADIIAAFYICII 1 2 gIMSATGNGYVIYMTIKRKSKLKPPELMTVNLAVFDFGIS 1 2 VTGKPFFVVSSFAHRWLFGWEGCRFYGWAGFFFGCGSLITMTVVSLDRYLKICHLRY 1 2 GTWLKRQHAFLCLVFVWMYAAFWATMPLVGWGNYAPEPFGTSCTLDWWLAQASVSGQSFVVAILFFCLVLPAGIIVFSYVMIIFKVKSSAKEISNFDARIKNSHNLEIKLTK 0 0 VAMLICAGFLIAWIPYAVVSVVSAFGEPDSVPISVSVIPTLLAKSSAMYNPIIYQVLDLKNSCMKSSCFKGLKKPRHFRKSR 2 1 YTISGSLKDTAPAKEAHIEI* 0 >NEUR1_pimPro 0 MENTSWPHSSYVPHYLLRGDPFASRLSKEADIVAAFYILII 1 2 GIMSATGNGYVIYMTIKRKSKLKPPELMTVNLAVFDFGIS 1 2 VTGKPFFVVSSFSHRWLFGWEGCRFYGWAGFFFGCGSLITMTVVSLDRYLKICHLRY 1 2 GTWLKRQHIFLCLVFVWIYAAFWATMPLVGWGSYAPEPFGTSCTLDWWLAQASVSGQSFVMSILFFCLVLPAGIIVFSYVMIICKVKSSSKEVSSFDARIKNSHTLEIKLTK 0 0 VAMLICAGFLIAWIPYAVVSVVSAFGEPDSIPIPVSVIPTLLAKSSAMYNPIIYQVIDCKKNCAKLSCFQAWSKRKHYKTSR 2 1 FYSISASMKKRPANEVPTEI* 0 >NEUR1_calMil 0 MTAFDNSTALYSGYWLHDSLHGDPFVSKLSWEADIISACYLIVT 1 2 GLLSTLGNGYVIYLSITQKRKLKPPEILITNLAISDFGMS 1 2 VGGQPFLIISCFSHRWIFGWVGCRWHGWAGFFFGCGSLITMTVVSLDRYLKICHLQY 1 2 GSWLQRRHVFMSLAFIWFYAAFWATMPLVGWGNYAPEPFGTSCTLDWWLARVSVSGLIFVLTILFFCLLLPIIIIVFSYIKIIAKVKSSAKEVAHFDSRIQNHHSLEMNLTK 0 0 2 1 * 0 >NEUR1_petMar Petromyzon marinus (lamprey) frag 2 GTWVRRRHAFLCVLAVWLYAAFWATMPLLGWGSYAPEPFGTSCTLDWWLAQSSAAGRSFVLCMLLFCLLLPAAAILFAYARIVGAVRRSARDLAHFERRARGGGGGGGGGGVALELRITK 0 0 VAMMICAGFLLAWIPYAVVSVWSAFGAPDSVPVAVSMVPTMFAKSAAMYNPLIYQLLSRRGTGAHCCRCRKARGTLRRPR 2 >NEUR1a_braFlo from cDNA and genome chrUn ++ 176419943 176431046 11104 FE548698 0 MATTPADRLDGLTPAGRGATTAETHADDFASKLSREADIVIGVYLILI 1 2 GTGAILGNGRVLWLSYRCRARLRPVEMFVVSLAAADVGLSLVGHPFSAASSLMGRWSFGSAGCTW 1 2 YGFVVFFLGIASIATMALMSIMRFMIVHKRY 1 2 GQYPSRRASCVLVAAAWLYGLFWACAPLA 1 2 GWSQYHPEPYGLSCSVDWGGFSRGAGGSSFIICMLLFCTAVPVVVMVTSYAAIFALYRQAQKGVVLNLQVNATFGGKRQRTER 0 IALAVCGGFLLAWLPYAVVGLWASVAGVDAVPLALASAAPLFAKSNSLWNPIIYLGMNERFR 2 1 * 0 >NEUR2b_braFlo from traces and genome chrUn ++ 187375671 187384042 8372 nearly identical chrUn ++ 32271780 32281075 9296 0 MATTPGLPLDGLAPTGRGVTAADTLDDDFASKLSREADIVIGVYLLLI 1 2 GTGSILGNGRVLWLSYRNWAKLRPVELFVVSLAVTDVGISVFGYPFAASSSLLGRWSFGSAGCTW 1 2 YGFTGFFFGLTSIANMALMSIMRFMIVYKGY 1 2 GPYPSRRATSGLIAAAWLYGLFWACAPLA 1 2 GWSQYHVEPFGLSCTVDWGSFSRDAGGMSFIICLLVFCVAIPVTAIMASYVAISAIYRQAKKSIAGHLQDNSAMCKKRNKLE 0 0 MALAVCGGFLLAWLPYAVVGLWSAVAGVDAVPLALASAAPLFAKSSSLWNPIIYLGMNDRFR 2 1 * 0 >NEUR1_strPur Strongylocentrotus purpuratus XM_001197837 CX694910 CX690664 0 MDVNAKWWTNETLRTRDQFSDDHYTSVLSYEGDIWAGVYLMFI 1 2 SLIAFIGNISVIVISLRKREKLKPIDLLTINLAIADFLICVVSYPLPMISAFRHR 0 0 WSFGKFGCVWYGFTSFLFAVGSMATLMVIALLRYAKLCRENV 1 2 DQYQSRPFVIKVIVAIWGFAFFTTAPPLFGWS 2 1 SYVPEPYHLSCTIDFADTSPSGLSYTYFTTIVVFFMPLMIIVLCYVAIARKMIHHNRRINVGHNAGRMLLEIRLLK 0 0 TACMITMAYTISWTPYAVIAMWVTYIPVNQIPDAFRILPAFCAKTSSVYNPIIYCIFNKSFRQDLSSLICCCACQCYTITINLDINSHAQQQFRRIEERR DEVGTYKRRPLMICSNPFAWSRDFHETWRQRRIRGIHRNCRNNVRVENINVNFRRDTDMVELNAPTPAEIHRPELNTASTRSGARTKSMATHLPALEEVPSG APQCSALLHNTPIPRSLQGTPLPYQPQPSTSDLHDEFLNPSVVSRNMCVIVVKPNIEEELSTD* 0
NEUR2: 11 vertebrate newropsins
>NEUR2_galGal Gallus gallus GenBank 5'UTR mistranslated as coding -B4GALT6 -NEUR2_galGal -KIAA1012 0 MDPSFANSTFQSKITEAADIVVGTCYMVF 1 2 GICSLCGNSILLYISYKKKHLLKPAEYFIINLAISDLAMTLTLYPLAVTSSLSHR 2 1 WLYGKHICLFYAFCGLFFGICSLSTLTLLSVVCCLKICFPAY 1 2 GNRFRRKHGQILIACAWTYAAIFACSPLAHWGEYGEEPYGTACCIDWQSTNVDVMSMSYTVVLFVLCFILPCGVIVTSYSLILVTVKESRKAVEQHVSGPTRINNVQTITAK 0 0 LSIAVCIGFFAAWSPYAIIAMWAAFGSIDKIPPLAFAIPAVFAKSSTLYNPIIHLLLKPNFRSNIAKDFTVIQQLCVRCCFCVKELQTYRSTFNTGLRTFKGKNESSCNALPIMEG CSYFPSEKGSHTFECFKSYPNCFQERLSTMGCHLQDCESLENDLQVEVTQGSRNSMKVVEQEEKSTELDNLEITLEAVPVSCTFTDL* 0 >NEUR2_anoCar Anolis carolinensis 0 MESYFANTTFHSKITEAADVIVGVFYIVF 1 2 GICSFCGNSILLYVSYKKKNLLKPAEYFMINLAISDLGMTLTLYPLAVTSSLAHR 2 1 WLFGQQVCLFYAFCGVFFGVCSLTTLTLLSIVCCLKICFPVY 1 1 GNRFRPGHGWILIACAWVYAAIFAFSPLAHWGEYGAEPYGTACCIDWRISNMKKTAMSYTTALFVFCYIIPCGIIITSYTLILITVKDSRKAVEQHALGPTRMSSVHTITAK 0 0 LSIAVCIGFFVAWSPYAIIAMWAAFGSIDMIPPLAFAVPAVFAKSSTLYNPAMYLFLKPNFRSTIAKDLTVLHRLCLKSCFCPRGMQNCSYRSALEAPLKSFKGRNESSSNSVQIVGGCS YFPCEKCHDPFECFKNYPKCCQGRLNVMDHTPRESISVENNMQSKTKHASEKYIKVVIRGEKNTDIDNLEITLEHIPTDIKFANL* 0 >NEUR2_xenTro Xenopus tropicalis abundant transcripts 0 MGNKSDASAFYSSISETDDIVLGVLYSVF 1 2 GLLSLSGNSMLLLVAYRKRSILKPAEFFIVNLSISDLGMTGTLFPLAIPSLFAHR 2 1 WLFDKVTCNYYAFCGMLFGLCSLTNLTVLSSVCCLKVCYPAY 1 2 GNKFSTAHSRILLLGIWAYAGLFATAPLADWGKYGPEPYGTACCLDWEASYRERKALSYTISLFVFCYLIPSSLIFISYTLIFVTVKGARRAVQQHLSPQAKGSSIHSLIIK 0 0 LSIAVCIGFLIAWTPYAIVAMMAAFGDPTKIPSLVFALAAAFAKSSTIYNPVVYLLLKPNFLNVVTKDLTLFQTMCAVVCGWCRTPAVKTPCPHKD LKTTSKPPSSFKKSQGVCRNCVDTFECFRNYPRCCSVGNVDAAQPMAASLVRIPPANGAPQQTVQLVVSSSRTRSGVETVEVSTEAPMSDFIKDFI* 0 >NEUR2_danRer Danio rerio acquired new intron 0 MGNVSKTALFMSTISRQHDILMGSLYSVF 1 2 FVLSLLGNGMLLFVAYRKRSSLKPAEFFVVNLSVSDLGMTLSLFPLAIPSALAHR 2 1 WLFGEITCLCYAVCGVLFGLCSLTNLTALSSVCCLKVCFPNY 1 2 GNKFSSSHACVMVIGVWCYASVFAVGPLVHWGSFGPEPYGTACCINW 2 1 YTPSHDALAMSYIISLFIFCYVVPCTIIILSYTFILVTVRGSQQAVQQHVSPQTKVTNAHALIVK 0 0 LSVAVCIGFLTAWSPYAIVAMWAAFSANEQVPPTAFALAAIMAKSSTIYNPMVYLLFKPNFRKSLSQDTQMFRHRICLSHSKASPSPGMKDQERQS SQQCNNKDGSISTPFSSGQAESYGACHVYAEAGPHYQQISRQITARVLEGSVQSEIPVKQLTEKMQNDLL* 0 >NEUR2_tetNig Tetraodon nigroviridis gene mix 0 MGNASDTSDAFNSKISKEHDFLIGSIYSVF 1 2 CVLSLMGNCILLLVAHHKRSTLKPAEFFIVNLSISDLGMTLTLFPLAIPSSFSHR 2 1 WLFGEIACQLYATCGVLFGLCSLTNLTVLSSVCCLKVCLPNL 1 2 GSKFSSSHARLLVAGVWGYASVFAVGPLVQWGHYGPEPYGTACCINWQAPNHELSSLSYIVCLFLFCYVLPCAIIILSYTCILMTVRGSRQAIQQHVSPQTKTANAHALIVK 0 0 LSVAVCIGFLGAWSPYAVVAMWASFGDATWVPPDAFAIAAILAKSSTIYNPLVYLLCKPNFRECLYKDTSTLRQRIYRGSPLSGPRDRSGGVTQRHKDLSVSTR LSNGQQDSYGTCLHCAEDAELGHVTGSRRTACILTGSTFTEVTLSQLSATPADLL* 0 >NEUR2_takRub Takifugu rubripes (fugu) 0 MGNASEASDIFLSKISKEHDILIGSIYSVF 1 2 GLLSLAGNCILLLVAYHKRSMLKPAEFFIINLSISDLGMTLTLFPLAIPSSFSHR 2 1 WLFGEITCQLYAMCGVLFGLCSLTNLTALSLVCCLKVCFPNH 1 2 GSRFSSSHARLLVVGVWCYASVFAVGPLVQWGHYGPEPYGTACCIDWRAPNHELSSLSYIVCLFFFCYVLPCATIILSYTCILMTVRGSRQAIQQHVSPQTKTANAHSLIVK 0 0 LSVAVCIGFLGAWSPYAIVAMWAAFGDATWVPPDAFAIAAILAKSSTIYNPVVYLLCKPNFRECLYKDTSTLRQRIYRGSPQSEPRERFGGTSQRHKDLSISTR LSNGQQDSYGTCLHCADDAERGHVTTSQRTACILTGSTFTEVTVGQLSAAPADLL* >NEUR2_gasAcu Gasterosteus aculeatus 0 MGNASDTSAVFASTISKERDILMGSLYSVF 1 2 GVLSLVGNCILLLVAYHKRSTLKPAEFFIINLSISDLGMTLSLFPLAIPSAFKHR 2 1 WLFGELTCQLYAMCGVLFGLCSLTNLTALSFVCCLKVCFPNH 1 2 GNRFSSSHARLLVVAVWGYASVFAVGPLARWGRYSPEPYGTACCIDWHAPNHELAALSYIVCLFVFCYALPCATIFLSYTFILLTVRGSRQAVQQHVSPQTKTTNTHALIVK 0 0 LSVAVCIGFLGAWTPYAVVAIWAAFGDATLVPPDAFALAAMFAKSSTIYNPVVYLLCKPNFRACLYRDTTLLRQRIYRGSPRSEPKAHFGSTSQRNKDMSVSVRSSNGQQDSYGACTENA APCHVMTPQRTACILTESTNREVTVSRLADKPQADFL* >NEUR2_oryLat Oryzias latipes 0 MGNVSDTSSLFASSISREHDILMGSLYSVF 1 2 GLLSLSGNSMLLLVAYRKRSILKPAEFFIVNLSISDLGMTGTLFPLAIPSLFAHR 2 1 WLFGEITCQLYAMCGVLFGLSSLTNLTALSLVCCLKVCFPNH 1 2 GNKFSFSHARLLVAGVWCYASVFAVGPLARWGRYSAEPYGTACCIDWHAPNHELWALSYILCLFIFCYALPCTIIFLSYAFILLTVRGSRQAVQQHVSPQTKTTNAHTLIVK 0 0 LSVAVCIGFLGAWTPYAVIAMWAAFGDATQVPPTAFALAAVFAKSSTIYNPMVYLLCKPNFRECLCRDTSLLRHMIYRGSPQPQERFGSDSRRNKDITASTRFSNGQQESYGACLNCTEN TGLCQLASPQNTACILTGSTYAEVTVQQLVDKQQPDFL* 0 >NEUR2_pimPro Pimephales promelas 0 MGNVSETALFVSTISRQHDILMGSLYSVF 1 2 CVLSLLGNGMLLFVAYRKRSSLKPAEFFVINLSVSDLGMTLSLFPLAIPSALAHR 2 1 WLFGEVVCLCYAVCGVLFGLCSLTNLTALSSVCCLKVCCPNY 1 2 GNKFSSNHACVMVIGVWCYASVFAVGPLIRWGSFAPEPYGTACCINWYIPSHDALAMSYIISLFIFCYVVPCTIIILSYTFILLRVRGSRQAVQKHVSPKTKETNAHTLIVK 0 0 LSVAVCIGFVTAWSPYAVVAMWAAFSANEPVPPTAFALAAILAKSSTIYNPMVYLLFKPNFRKILSQDTQNIRHRMCVSHSKASPTPEIK-AQSSQQCKDATISTPFSSGQAESYGTCHIYAEAEPHFQQISPQRTVRILEGIIQSEISVRHMTDRMQNDLL* 0 >NEUR2_oncMyk Oncorhynchus mykiss no glycosylation site, anomalous agreement with chicken 0 MGVLASIDDIAFLSNIPVAADITVAIVYAVF 1 2 GMCSLFSNSTLLYISYKKKHLLKPAEFFIINLAISDMSLTLSLYPMAITSSIYHR 2 1 WLFGKTVCLIYAFCGMLFGVCSLTTLTLLSMVCFVKVCYPLY 1 2 GNRFNAVHGRLLIACAWAWALVFACSPLAHWGEYGPEPYGTACCIDWRLSNLHPVARSYTAALFVLCYIVPCCVIVASYTGILMTVRASHKAMEHHEARQTKMSNIQDVIVK 0 0 LSVAVCIGFFAAWSPYAVVSMWAAFGHMDNIPPLAFAVPAMFAKSSTIYNPIIYLLLRPNFRRVMYRDLVSLCRAFLKGCLCSCSQGAVGKCHSHLVVRVSLQSFCRLPGHGQ SCSPTSSARQALGESRGCTSPGEKCSDAFECFRHYPRGCHGGTNIPSSSARVYAPQDQLSTEPQLQSMTQKQMRKQEACHKKSLRATKHSKRTSEIDNLRINFEMVPGHAKVAWP* 0 >NEUR2_calMil frag 0 1 2 GILSLVGNSVLLFVAYRKRQILKPAEYFVANLAVSDISMTVTLLPLAISSNFSHR 2 1 WLFVSKpCMYYGFCSMLFGICSLTNLTVLSTVCCMKVCFPAY 1 2 0 0 MSVVMIVMFLLAWSPYSIVCLWASFGNPKLIPPAMAIIAPLFAKSSTFYNPCIYVISYTMTVIAVNFVVPLSVMFFCYYNV
NEUR3: 16 vertebrate newtopsins
>NEUR3_galGal Gallus gallus cOpn5L2 mRMA for Opsin 5-like 2 AB368183 chr3 XM_420056 CN231992 testis exon 2^3 rel NEUR1/2 0 MEEQYISKLHPVVDYGAGVFLLII 1 2 AILTILGNSAVLATAVKRSSLLKSPELLTVNLAVADIGMAISMYPLAIASAWNHAWLGGDASCIYYALMGFLFGVCSMMTLCAMAVIRFLVTNSSKSN 1 2 SNKISKNTVHILITFIWLYSLLWAILPLVGWGYYGPEPFGISCTIAWSKFHSSSNGFSFILSMFLLCTVLPALTIVACYLGIAWKVHKAYQEIQNINRIPHAAKLEKKLTL 0 0 MAVLISVGFLSAWTPYAAASFWSIFNSSDSLQPIVTLLPCLFAKSSTAYNPFIYYIFSKTFRHEIKQLQCCWGWRVHFFSADNSAENSVSMMWSGRDNIRLSPTAKVESQGAARH* >NEUR3_taeGut Taeniopygia guttata ABQF01025032 0 MEEQYISKLHPVVDYGAGVFLLII 1 2 AILTILGNSAVLATAVKRSSLLKPPELLTVNLAVADIGMALSMYPLAIASAWSHAWLGGDASCVYYALMGFLLGVCSMMTLCAMAVIRFLVTNSPKSN 1 2 sNKITKNTVCILIAFIWLYSLLWAILPLVGWGYYGPEPFGISCTIAWSKFHNSSNGFSFILSMFLLCTVLPALTIVACYLGIAWKVHKAYQEIQNIDRIPNAAKLEKKLTL 0 0 MAVLISVGFLSSWTPYAATSFWSIFNSSHSLQPVVTLLPCLFAKSSTAYNPFIYYVFSKTFRCEVKRLQCCCAWRVHYFSSDNSVENPLSTMWSGRDNIRLSAAPQVQNPGAAAP* 0 >NEUR3_anoCar Anolis carolinensis AAWZ01001057 0 MEEHYISKVHPVWDYGMGVFLLII 1 2 AILTILGNSMVLAVAVKRSSCLRSPELLTVNLAATDLGMGLSMYPLAIASAWNHAWLGGEATCIYYALMGFLFGVSSIMTLSAMAVIRFLVTFSSKPA 1 2 GHKINRKVMHICIMLIWAYAVLWAILPLLGWGHYGPEPFGTSCTIAWGQFHNSQKGFAFILSMFILCTFLPAITIIMCYLGIAWKFHKTHQEMQNLNRISSAAKLEKKLIL 0 0 VAVLISVGFLGAWTPYAIVSFWSVFHSSESIPYIVTLLPCLFAKSSTAYNPFIYYTFSKTFRHEVKHLRCYSGQRAQENMKNSINSNVSFMWHGGGNICLSTRQIEMREIPNQ* 0 >NEUR3_xenTro Xenopus tropicalis cdna ovary embryo 0 MEERYLSKLHPLVDFGSGVFLLLV 1 2 AILTVLGNCAVLATAVKCSSHLKAPDLLSINLAVADLGMAISMYPLAIASAWNHAWLGGDASCLYYALMGFFFGVSSMMTLTVMAIIRYRVTSSFKYS 1 2 GCTIEKKAVCILIMCIWLYALLWAVLPLLGWGRYGPEPFGTSCTIAWGDFHHSSNGFSFIISMFILCTISPAVTIVVCYSGIAWKLHKAYQEIKNQDKIPNSTKVEKKLTL 0 0 LAILVSFGFLISWTPYAAVSFWSLFHSSKYIPPVVSLLPCLFAKSSTAFNPMIYYAFSKTFRRKVKHLKCCCGWRVHFLQSENSVENPRVSVIWTGKENVMVSSVPKLMKGVPGTPTGTQ* 0 >NEUR3a_danRer Danio rerio 0 MDRYTSKLSPAVDYSAGTFLLVI 1 2 AILSILGNAAVLLTAAWRHSVLKAPELLTVNLAVTDIGMALSMYPLSIASAFNHAWIGGDPSCLYYGLMGMIFSVASIMTLAVMGLVRYLVTGNPPK 1 2 SGSKFRRKTISILIGVIWMYSLLWAVFPILGWGGYGPEPFGLACSVDWMGYQHSLNRSSFIMALAILCTLMPCVVILFSYSGIAWKLHKAYQSIQSNDNLPNSGAVERKVTL 0 0 MGILISTGFIVSWAPYVFVSLWTMFRSEGEDSVVPIVSLLPCLFAKCSTVYNPLVYYVFRKSFRREIHQIRICCFQGCWDAVSKMTRGDGPEETSGTHETDNI* 0 >NEUR3a_tetNig Tetraodon nigroviridis 0 MDDKYMSKLSPPVDLWAGIYLVVI 1 2 ALLSVLGNASVLFSASRRLTPLKAPELLTVNLAVTDIGMALSMYPLSIASAFNHAWMGGDTACLYYGLMGMIFSITSIMTLAVMGMIRYLVTGSPPR 1 2 SGVQFQKKTICVVICAIWLYACLWAAFPLLGWGSYGPEPFGTACSIDWTGYGDSLNNATFIVAMSVLCTFLPCLVIFFTYFGIAWKLHKAYKSIKSSDFQYASVERRITL 0 0 IAVLISVGFLGSWAPYGLVSLWSILKDSSSIPPQVSLLPCLFAKSSTVYNPVIYYIFSQSFKLEVQQLFLCC* 0 >NEUR3a_takRub Takifugu rubripes (fugu) 0 MDDKFTSKLSPAVDLWAGTYLVFI 1 2 ALLSVLGNASVLFSAGRRLSMLKAPELLTVNLAVTDIGMALSMYPLSIASAFNHAWMGGDASCLYYGLMGMIFSITSIMTLAVMGMIRFLVTGTPPR 1 2 SGIKFQKKTISVVISAIWLYACLWAVFPILGWGSYGPEPFGIACSVDWMGYGESLNNATFISTLSVLCTFLPYLVIVFTYFGIAWKLHRAYRSIKSSDIQYTNVERRITL 0 0 MAVMISSGFLIAWTPYVAVSFWSMRNSQRQGHMAPSVTLLPCLFAKSSTAYNPFIYFFFQRNTGHKLLPFHRHAFSCSDRADSSREGEKEESKVSKNLGFTCFGAGTYETCPGLAGDQSQREMAELG* 0 >NEUR3a_gasAcu Gasterosteus aculeatus 0 MEDKYVSKLSPAVDFWAGTYLIII 1 2 AVLSIFGNTAILVSAARRSGPLKAPELLTVNLAVTDLGMALSMYPLSIASAFNHAWIGGDASCLYYSLMGMIFSITSIVTLAVMGMVRYLVTGNPPR 1 2 SGLRLQRKTVSMVIGAVWLYSGLWALFPLLGWGSYGPEPFGLACSIDWSSYGESLNRSTFIMTLSVLCTFLPCLVIFFTYFGIAWKLRRAYQSIRSSDFQHGKVEQKITL 0 0 MAAMISSGFLFSWTPYVAVSLWSMFRSREHIPPLVALLPCLFAKSSTVYNPFIYFIFQRSSWRELLRLHRHLLCCWHRASPPAEGRRSQRGSEGGSWGGACESDDAFGLVHVMKSNATCQTISWA* 0 >NEUR3_calMil frag 2 AILSIFGNSVVLLVAAKKSSQLKPPELLTVNLAITDFCSAVTMYPLAVGSAWKHTWLGGDASCKYYGFMDFFFGIASIGTLTVMAIVRFLVTSTTQN 1 >NEUR3_petMar new exon frag 0 MAEQGEDDQFRSKLSPTADIAAGTFLLAV 1 2 AVLSLAGNGAVLGVAARRWAKLKAPELLSVNLALTDLGIAASIYPLAVASAWNHRWLGGQPVCTYYAFAGFFFGTASMGTLTAMAGVRYKGTSTQVH 1 2 sVKQITKRAMLAVIVAVWAYALLWSCLPLLGWGR 2 1 YGVEPFGVSCTLAWAELQLTPGGVAFLYAMFVLCLLLPAIAIGLCYAGIVCKLRRAYREGRSKRRTPTARHVESRLTK 0 >NEUR3b_danRer Danio rerio 0 MDIYSSKLSSAVDYGIGAFLLLI 1 2 TILSILGNLMVLVMAYKRSNHMKPPELLSVNLAVTDLGAAVTMYPLAVASAWNHHWIGGDVSCVYYGLMGFLFGAASMMTLTIMAIVRFIVSLTLQSP 1 2 KEKISKRNAKILVATTWLYALLWAIFPLIGWGKYGPEPFGLSCTLDWRDMKEHSQSFVITIFLMNLILPAIIIVSCYCGIALRLYVTYKSMDDSNHVPNMIKMQRRLMV 0 0 IAVLISIGFVGCWAPYGIVSLWSIYRPGDSIPAEVSMLPCLFAKTSTVYNPFIYYIFSKTFKREVNQLSRFCGRSNICRPTDAKNRPENTIYLVCDVNKSKPGVEDLSLARSKENETQMLPNQDLHE* 0 >NEUR3b_tetNig assembly errs in exon 2 frameshift, used traces 0 MDMYTSALSPALDIGTGCYLLVI 1 2 AVLSFIGNLLVIITAVKKSSKMKPPELLCVNLAVTDLGAAVTMYPLSVASAWSHRWIGGDVTCVYYGLVGFLFEVASIMNLTVLAIVRFTVSLNLQSP 1 2 EEKISWKSVKIMCLLIWLYGVIWAMFPVLGWGRYGPEPFGISCSLAWGQMKNEGFSFVVAMFSFNLAVPALIIVSCYFGMAINLYFTHKKMVNTGNRIPAVIKLHRRLLR 0 0 IAVLISVGFLGSWAPYGLVSLWSILKDSSSIPPQVSLLPCLFAKSSTVYNPVIYYIFSQSFKLEVQQLFLCCLSFRSSRTNNCKSNESSIFMVSNGKNLTPALTQQNTSHAVIMN* 0 >NEUR3b_takRub 0 MDIYSSTLSPALDIGTGCFILVV 1 2 GVLSIIGNLLVIITAVKRSSKMKPPELLCVNLAVTDLGAAVTMYPLSVASAWSHRWIGGDATCIYYGLVGFLFGVASIMNLTILAIVRFTVSLNLQSP 1 2 eEKITWKSVKIMCMWVWLYSIMWAMFPILGWGRYGPEPFGISCSLAWGQMKDEGFSFVVTIFSLNFAVPAVIIICCYFGIAIKLYFTYKKTVNTNQIPVIIKLHRRLLM 0 0 IAVLISVGFLGCWAPYGLVSLWSILKDSSSIPPEVSLLPCMFAKSSTVYNPIIYYMFSQSFKMEVQQLFLWCPSFEFCRTSSNNGNETTIYMVSTGKT* 0 >NEUR3b_gasAcu 0 MDIYASTLSPAVDVGAGCYLLFV 1 2 AVFSIVGNLLVLVMAVKRSSRMKPPELLSVNLAVTDLGAAVTMYPLAVASAWRHRWLGGDATCVYYAVAGFFFGLASIMSLTGLAIVRFIVSLNLQSP 1 2 NEKISWRKVKLLCACTWLYALAWAAFPFLGWGRYGPEPYGLSCSLAWGQMKHEGFSFVVSMFSLNLVLPCVIIAGCYFGIAFKLYFTYRKSNNNSNRLPNVVRRHRRLLA 0 0 IAVLISLGFVVCWSPYAVVSLWSIFHDSGSIPPEVSLLPCMFAKSSTVYNPLIYYIFSQSFRREVKQLWRHLGSTLCSVSNSVNDAAVSNTGKSN* 0 >NEUR3b_oryLat 0 MDIYASALSPALDIGTGCYLLVL 1 2 TVLSIIGNLLVVIMAFKRSSRMKPPELLSVNLALTDLGAAVFMYPLAVASAWSHHWLGGDVSCIYYGLAGFFFGSASVMNLTALAVVRFIVSLNLHSP 1 2 KEKVSWRKVKILCLWSWLYALIWALFPILGWGRYGPEPFGLSCSLAWGEMKQEGPSFVISLFSFNLVLPSVVIICCYFGIAMKLYFTYKKSANSNHVPNIIKLHRRLLIIA 0 0 ILISIGFIGCWTPYGLVSLWSIFNDSSKIPPEVSLLPCMFAKSSTVYNPMIYYFFSKSFQREVKQLSWLCVGSNPCHVSNSVNDNNIYMVSVNVKSKETRRETLQEITESRQ* 0
NEUR4: 10 vertebrate newwopsins
>NEUR4_ornAna Ornithorhynchus anatinus hypothetical protein XM_001508128 0 MSLSHSLQVPWRNNLTFLNKEAQVSEQGETIIGIYLLAL 1 2 GWMSWFGNSMVIFILHRQRGILNPTDYLTFNLAVSDASVSVFGYSRGIIEIFNVFRDDGFLITSIWTCQ 0 0 VDGFLTLLFGLASINTLAMISVTRYIKGCHPHR 1 2 GHFINTANISVALILIWVSALFWSAGPVLGWGSYT 1 2 DRMYGTCEIDWAEANFSSICKSYIISIFFCCFFLPVSIMFFSYVSIIKMVKSSHTLAGADDPTDRQRRLDRDVTR 0 0 VSVVICTAFIVAWSPYAVISMWSAFGHSVPNLTSVLASLFAKSASFYNPIIYFGMNSKFRKDILVLLPCAKESKEPVKLKKFKNLRQKQGFTLQKPEKAHVLQVPDSGPMSLINTPPLGNRNSFDLACDNSDFECVRL* 0 >NEUR4_galGal Gallus gallus genome gappy 0 MSLQLSPQAPWRNNNISFLSREAAVTEQGETIIGFYLLAL 1 2 GWMSWFGNSVVIFVLYKQRHLLQPTDYLTFNLAVSDASISVFGYSRGIIEIFNVFRDDGFIITSIWTCQ 0 0 VDGFLTLLFGLASINTLTVISVTRYIKGCHPER 1 2 AHCISNSSMTVAMVLIWIAAFFWSAAPLLGWGSYT 1 2 DRMYGTCEIDWAKANFSTIYKSYIISIFICCFFLPVTVMVFSYVSIINTVKLSHALTGLSDPTERQRRMERDVTR 0 0 IVICTAFIIAWSPYAVLLLWSAYGHPVPNLPLYLSSLFAKSASFYNPIIYFGMSSKFRRDIFILFHCAKEVKDPVKLKRFKNLKQKQEPSQKEEKYAAEMHPAPSPDSGVGSPTNTPPPANREEYFGILDTPSNSPDIECDRL* 0 >NEUR4_taeGut Taeniopygia guttata 0 MSVQFSAQAPWRNNNISFLTREAAVTEQGETIIGFYLLAL 1 2 GWLSWFGNSIVIFVLYKQRHVLQPTDYLTFNLAVSDASISVFGYSRGIIEIFNVFRDDGFIITSIWTCQ 0 0 VDGFLTLLFGLASINTLTVISVTRYIKGCHPER 1 2 GHCISNSSMSVALVLIWVAAFFWSAAPLLGWGSYT 1 2 DRMYGTCEIDWAKASFSTIYKSYIVSIFICCFFLPVTVMVFSYVSIINTVKLSHT LTGLGDPTDRQRRIERDVTR 0 0 VSLCTAFIIAWSPYAVISIWSAYGHPVPNLTSILASLFAKSASFYNPIIYFGMSSKFRRDIFIFHCAKELKDPVKLKRFKNLKPKQPQPSQKEEKYAPEMHPAPSPDSGVGSPTNSPPPANREVYFGILDTPSNNPNIECDRL* 0 >NEUR4_anocar Anolis carolinensis 0 MSLQVSPQAPWRNNNVTFSNKEVPVSEQGETIIGFYLLAL 1 2 GWMSWFGNSIVIFVLYRQRAGLQPTDYLTFNLAVSDASVSVFGYSRGIIEIFNVFRDDGFLITSIWTCQ 0 0 VDGFLTLLFGLASINTLTVISVTRYIKGCHPDR 1 2 GKCISNSSISVALFLIWIAAFFWSVAPVLGWGSYr 1 2 DRMYGTCEIDWAKANFSTIYKSYIVSIFICCFFLPVSVMVFSYVSIINTVKSSHALSGVGDPTERQRRMERSVTR 0 0 VSLVVICTAFITAWSPYAVISMWSAYGYTVPNLTSILASLFAKSASFYNPIIYFGMSSKFRKDIFVLLHCAKEIKDPVKLKRFKNLKQKQEVSPSQREEKYAADVQPALSPDSGVGRSNTPPPVNREVYFGAFDTFSNNPDVECDRL* 0 >NEUR4_xenTro Xenopus tropicalis 38% NEUR1_galGal 0 MSLQFPRPAPWRNNNLTLLQKENPLTEQGETIIGIYLLAL 1 2 GWLSWFGNSIVIFVLYKQRANLLPTDYLTFNLAVSDASTSVFGYSRGIIEIFNVFRDDGFLITSIWTCQ 0 0 VDGFLTLLFGLASINTLTLISVTRYIKGCHPQR 1 2 ANCISNGSITISLALIWIAALFWSVAPLLGWGSYR 1 2 DRMYGTCEIDWTKASFSTIYKSYIISIFICCFFLPVMVMVFCYVSIINTVKSSRALTSEGDLSERQRKMERDVTR 0 0 VSVVICTAFIVAWSPYAVISMWSACGYYVPSLTSILAALFAKSASFYNPLIYFGMSSKFRKDLCVVLPCAKAQKDPVKLKRYKDKKQGSAPRAREQTEIEQPVQLQPAPSQDSGVGSPSNTPPLRTKDVHIVDIDLVSDNPSYECDRL* 0 >NEUR4_danRer Danio rerio 0 MSAQNPLQVVNIPWRNNNFSLMSRDPPLSDQGETIIGVYLLIL 1 2 GWLSWFGNSIVIFVLFRQRSTLQPTDYLTLNLAVSDASISVFGYSRGILEIFNIFKDSGYIISSVWTCQ 0 0 VDGFFTLVFGLSSINTLTVISITRFIKGCHPHK 1 2 AHCITNSTVAVCVVFIWIGAFFWSAAPVLGWGSYT 1 2 DRGYGTCEIDWVKANYSTIHKSYIISIFIFCFLVPVLLMLFCYISIINTVKRGNAMNADGDLSDRQRKIERDVTI 0 0 VSIVICTAFILAWSPYAVVSMWSAWGFHVPNLTSIFTRLFAKSASFYNPLIYFGLSSKFRKDVSVLLPCGREGRDPVRLKRFKRLRGRAEPPGAPAHTPHPQIALKNYNNHSKPHAGPAHCTGHAPSPDSGVGSHHETPPPQPRPQLFFIDVPEPEAESECVRL* 0 >NEUR4_tetNig Tetraodon nigroviridis 0 MEPSRPWRNSSVLGGGAEPPLSEQGETIIGVYLLLL 1 2 GWLSWFGNTVVLFVLVRQRSSLQPTDLLTFNLAVSDASISVFGYSRGIIQIFNVFQDSGFIISSIWTCE 0 0 VDGFLTLIFGLSSINTLTVISITRYIKGCQPSR 1 2 AALISRSSVSVCLLLIWTTAGFWSGAPLLGWGSYT 1 2 DRGYGTCEIDWSKAASSGVYRSYIISIFIFCFFIPVFIMLFCYISIINTVKRGNALAADGHLSHRQRTMERDVTV 0 0 ISVVICTAFIMAWSPYAVVSMWSAWGFHVPSTTSIVTRLFAKSASFYNPLIYFGMSSKFRKDVSLILPCAKERREVVLLQRFKNIKPKAAAAPPPPPLPVYRPKEKNEDEPKLSVHDNDSGVNSPPETPPSDAQEVFPVDPPSQIETSEYWSDRL* 0 >NEUR4_takRub recent pseudogene 8/8 traces support stop codon indel too 94% identity 0 MADSIPPWRNSSVLGGGAEPPLSEQGETIIGVYLLLLG 1 2 GWLSWFGNTVVLFVLYRQRSTLQPTDYLTFNLAVSDASISVFGYSRGIIEIFNVF*DSGFIISSIWTCE 0 0 VDGFFTLVFGLSSINTLTVISITRYIKGCQPSR 1 2 AGHINRTFVSVCLLLIWIMAGFWSGSPLLGWGSYT 1 2 DRGYGTCEIDWSKAAYSTAYRSYIISIFIFCYFIPVFIMLFCYISIINRVKRGNALAA-GDLTDRQRKMERDVTI 0 0 VSIVICTAFILAWSPYAVVSMWSAWGFHVPNLTSIFTRLFAKSASFYNPLIYFGLSSKFRKDVAVLLPCTKDAKDTVKVKRFK NIKPKAAAAPPPPPLPVYRPKEKNEDEPKLSVHDNDSGVNSPPETPPSDAQEVFPVDPPSQIETSEYWSDRL* 0 >NEUR4_gasAcu Gasterosteus aculeatus (stickleback) 0 PVKVVNIPWRNNNLSNLNTDPPLSEQGETFIGVYLLVL 1 2 GWLSWFGNSLVMFVLYRQRASLQSTDFLTLNLAISDASISIFGYSRGILEIFNIFNDDGYLINWIWTCQ 0 0 VDGFFTLLFGLASINTLTVISVTRYIKGCHPNK 1 2 AYCISTNTIAVSLICIWTGAVFWSVAPLLGWGSFT 1 2 DRGYGTCEVDWSKANYSTIHKSYIISILIFCFFIPVMIMLFSYVSIINTVKSTNAMSADGFLSTRQRKVERDVTRV 0 0 ISIVICTAFITAWSPYAVVSMWSAWGFHVPSTTSIITRLFAKSASFYNPLIYFGMSSKFRKDVSVLVPCTRERREVVHLQHFKNIKPKAEAPPTPASLPVQKLGAKYAVPNPDADSGVNNPPQRPATDPQGDLNIDLPSHIETSEYWCDRL* 0 >NEUR4_oryLat Oryzias latipes (medaka) frag 0 MEITLKAFPLKVVNIPWRNNNLSTLHSEPPLSEQGETVIGVYLLVL 1 2 GWLSWFGNSLVIFVLCKQRASLQPTDFFTLNLAVSDASISVFGYSRGILEIFNILKDDGYLITWIWTCQ 0 0 VDGFLTLLFGLVSINTLTVISVTRYIKGCHPHK 1 2 AHCISSSTIAVSLIIVWAAALFWSVAPLLGWGSYT 1 2 DRGYGTCEVDWSKANYSTFYKSYIISILIFCFFIPVVIMLFSYVSIINTVKSTNAMSAVGFLSARQRKMERDVTRV 0 0 ISIVICTAFITAWSPYAVVSMWSAWGFHVPSTTSIITRLFAKSASFYNPLIYFGMSSKFRKDVSVLVPCTRERREVVHLQHFKNIKPKAEAPPTPASLPVQKLGAKYAVPNPDADSGVNNPPQRPATDPQGDLNIDLPSHIETSEYWCDRL* 0 0 SASFYNPLIYFGMSSKFRKDISVLLPCAAEGREVVHLQRFQNIKPKADTPLTAAPHPPPAKPLAAEMNQTNADGDPGVNNPPHTPPQIFHIDVPSHIETSEFWCDRL* 0 >NEUR4_calMil Callorhinchus milii frag 0 MGCSLGWKVLLWFLHGILICPRPWRNHNSTFQPKEHPISEQGETIIGVYLLIL 1 2 GWLSWFGNSIVIFILYRQRLSLQPPDYLTLNLAVSDASISIFGYSRGIIEIFNVFRDDGFLITSIWTCQ 0 0 1 2 AVSISAGSIAASLVLIWIAAIFWSGAPLFNWGSYT 1 2 DRMYGTCEIDWSRASFSTIYKSYIISIFICCFFLPVFVMLFSYISIINTVKSSHAFAGNADLSDRQRRMEKDVTR 0 0 VSMVICTAFIIAWSPYAVISMWSASGYTVPQLTGIFASLFAKSASFYNPMIYFGLNSKFRKDIYILLPCVKEPKESVKLKRFKHLRHRPEQQQANKDRYAEELQQVASPDSGMGSPSKSPPLHNKDVFFVLWLRGLKK >NEUR4_petMar lamprey frag 0 1 2 GWLSWLGNGLVIFVLTRQWSSLQPPDLLTLNLALSDASIAVFGYSRGIIEIFNVFQDDGYIIKSTWTCQ 0 0 1 2 PTKVTSTSMVVSLALVWAASLFWSAAPLLGWGSYT 1 2 DRRYGTCEIDWMKATFSTIYKSYIISIFICCFFMPISTMLFAYISIINTVKSSHVTARMGDVSERQRNMERDITRI 0 0 VSIVICCAFILAWSPYAVISMYSACGHRVPALTSLLAALFAKSASFYNPFIYFGMSGKFRADVRAMLPCRATSVKAPRDAVRLKRYRTHVDPERASHRAAVAAREQPAPRAAAPRPASPAPSAARDRDPELDEREFDPEGRASALAEVAAVESRDSGIACTRGKRRASRGDDVEVRNDV* 0
See also: Curated Sequences | Peropsins | RGR phyloSNPs | LWS | Encephalopsins | Melanopsins | Update Blog
