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Evolution of the Relaxin Insulin Like Gene Family in Anthropoid Primates Jose Ignacio Arroyo1 2 Federico G Hoffmann3 4 andJuanC Opazo1 1Instituto de Ciencias Ambientales y Evolutivas

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	EvolutionoftheRelaxin/Insulin-LikeGeneFamilyin
AnthropoidPrimates
IgnacioArroyo
,FedericoG.Hoffmann
,andJuanC.Opazo
InstitutodeCienciasAmbientalesyEvolutivas,FacultaddeCiencias,UniversidadAustraldeChile,Valdivia,Chile
ProgramadeDoctoradoenCienciasmencio
nEcolog
ayEvolucio
n,FacultaddeCiencias,UniversidadAustraldeChile
DepartmentofBiochemistry,MolecularBiology,EntomologyandPlantPathology,MississippiStateUniversity
InstituteforGenomics,BiocomputingandBiotechnology,MississippiStateUniversity
*Correspondingauthor:E-mail:[email protected]
January29,2014
Therelaxin/insulin-likegenefamilyinclud
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Thenumberandnatureofgenesinthesethreegenomic
lociarewellconservedinmostmammalianlineages,withthe
exceptionoftheRFLBlocus(
Parketal.2008
Hoffmannand
Opazo2011
;
Arroyo,Hoffmann,Good,etal.2012
Hoffmann,Opazo2012a
2012b
).Thislocuspossessacom-
plexduplicativehistorycharacterizedbysmall-scaleduplica-
ABC
.1.
492
GenomeBiol.Evol.
6(3):491–499.doi:10.1093/gbe/evu023AdvanceAccesspublicationFebruary2,2014
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EvolutionofRLN1andRLN2GenesinAnthropoidPrimates
GenomeBiol.Evol.
6(3):491–499.doi:10.1093/gbe/evu023AdvanceAccesspublicationFebruary2,2014
493
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proposedtoexplainthetimingofduplicationandgenegains
andlossesalongtheorganismaltree(
g.1
).Initialstudies
hadsuggestedthattheduplicationgivingrisetoRLN1and
494
GenomeBiol.Evol.
6(3):491–499.doi:10.1093/gbe/evu023AdvanceAccesspublicationFebruary2,2014
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.2.
Maximumlikelihoodphylogramsdepictingrelationships
amongrelaxin-likegenesinprimatesbasedon1kbof5
ankingsequence,codingsequence,intron1,and1kbof3
sequence.Numbersonthenodescorrespondtomaximumlikelihoodbootstraps
upportvaluesandBayesianposteriorprobabilities.SinglecopyRLNgen
efoundinNewWorldmonkeysareshaded.
EvolutionofRLN1andRLN2GenesinAnthropoidPrimates
GenomeBiol.Evol.
6(3):491–499.doi:10.1093/gbe/evu023AdvanceAccesspublicationFebruary2,2014
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astheLRTsfavoredthealternativemodel(LRT
0.049),whereseveralresiduesswitchedtoapositiveselec-
tionregimeintheancestralbranchoftheNewWorldmon-
keysRLNclade.TheBEBanalysisidentied35codonsundera
positiveselectionregime,twoontheregionencodingforthe
signalpeptide,fourontheregionencodingfortheBpeptide,
21ontheregionencodingfortheCpeptide,andeight
locatedontheregionencodingfortheApeptide(
table1
TheseresultssuggestthatpositiveDarwinianselectioninthe
ancestralbranchoftheNewWorldmonkeyRLNcladewas
responsiblefortheremodelingofthisprotein,andprobably
Table1
.3.
AnevolutionarymodelfortheevolutionoftheRLN1and
RLN2genesinanthropoidprimates.ThemodelindicatesthattheRLN1
andRLN2paralogsderivefromtheduplicationofaproto-RLNgeneinthe
lastcommonancestorofcatarrhineprimates,andnotinthelastcommon
ancestorofapesoranthropoidsaspreviouslythought.Althoughatwo
genearrangementwaspresentinthelastcommonancestorofcatarrhine
primates,onlyapesappeartohavere
tainedbothcopies,whereasOld
WorldmonkeyslosttheRLN1paralog.
496
GenomeBiol.Evol.
6(3):491–499.doi:10.1093/gbe/evu023AdvanceAccesspublicationFebruary2,2014
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.4.
Alignmentsofrelaxinaminoacidsequences.TheupperpaneldepictsanalignmentoftheancestralstatesreconstructedforthebranchleadingtotheN
ewWorldmonkeyRLNclade(nodes1
and2),andtwoactualNewWorldmonkeyspecies.Th
emiddlepanelshowsanalignmentoftheancestralstatesreconstructedforthebranchleadingtothe
apeRLN1clade(nodes3and4),andveactual
apespecies.ThelowerpanelshowsRLN2sequencesfromveactualapespec
ies.Aminoacidsinbolddenotesitesinferredunderpositiveselection,sha
dedaminoacidsareparallelchanges,andboxed
aminoacidisaparallelchangewherethederivedaminoacidstatewasnotth
esameinbothlineagesbuttheybelongtothesamefunctionalgroup.Aminoac
idsiteslabeledwithanXwerenotincludedin
theancestralsequencereconstructionanalysisastheentirecolumnsofgappedsiteswereremoved.
EvolutionofRLN1andRLN2GenesinAnthropoidPrimates
GenomeBiol.Evol.
6(3):491–499.doi:10.1093/gbe/evu023AdvanceAccesspublicationFebruary2,2014
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possessesbiologicalactivitysimilartotheprocessedpeptide
498
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6(3):491–499.doi:10.1093/gbe/evu023AdvanceAccesspublicationFebruary2,2014
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ArroyoJI,HoffmannFG,OpazoJC.2012b.Geneturnoveranddifferential
retentionintherelaxin/insulin-likegenefamilyinprimates.Mol
EvolutionofRLN1andRLN2GenesinAnthropoidPrimates
GenomeBiol.Evol.
6(3):491–499.doi:10.1093/gbe/evu023AdvanceAccesspublicationFebruary2,2014
499
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