title: A new model army: Emerging fish models to study the genomics of vertebrate Evo-Devo.
abstract: Many fields of biology--including vertebrate Evo-Devo research--are facing an explosion of genomic and transcriptomic sequence information and a multitude of fish species are now swimming in this "genomic tsunami." Here, we first give an overview of recent developments in sequencing fish genomes and transcriptomes that identify properties of fish genomes requiring particular attention and propose strategies to overcome common challenges in fish genomics. We suggest that the generation of chromosome-level genome assemblies--for which we introduce the term "chromonome"--should be a key component of genomic investigations in fish because they enable large-scale conserved synteny analyses that inform orthology detection, a process critical for connectivity of genomes. Orthology calls in vertebrates, especially in teleost fish, are complicated by divergent evolution of gene repertoires and functions following two rounds of genome duplication in the ancestor of vertebrates and a third round at the base of teleost fish. Second, using examples of spotted gar, basal teleosts, zebrafish-related cyprinids, cavefish, livebearers, icefish, and lobefin fish, we illustrate how next generation sequencing technologies liberate emerging fish systems from genomic ignorance and transform them into a new model army to answer longstanding questions on the genomic and developmental basis of their biodiversity. Finally, we discuss recent progress in the genetic toolbox for the major fish models for functional analysis, zebrafish, and medaka, that can be transferred to many other fish species to study in vivo the functional effect of evolutionary genomic change as Evo-Devo research enters the postgenomic era.
authors.affiliation: Institute of Neuroscience, University of Oregon, Eugene, Oregon.
authors.forename: Ingo
authors.initials: I
authors.lastname: Braasch
authors.affiliation: Institute of Neuroscience, University of Oregon, Eugene, Oregon.
authors.forename: Samuel M
authors.initials: SM
authors.lastname: Peterson
authors.affiliation: Institute of Neuroscience, University of Oregon, Eugene, Oregon.
authors.forename: Thomas
authors.initials: T
authors.lastname: Desvignes
authors.affiliation: Institute of Neuroscience, University of Oregon, Eugene, Oregon.
authors.forename: Braedan M
authors.initials: BM
authors.lastname: McCluskey
authors.affiliation: Institute of Neuroscience, University of Oregon, Eugene, Oregon.
authors.forename: Peter
authors.initials: P
authors.lastname: Batzel
authors.affiliation: Institute of Neuroscience, University of Oregon, Eugene, Oregon.
authors.forename: John H
authors.initials: JH
authors.lastname: Postlethwait
mesh_terms: D000818:Animals; D019143:Evolution, Molecular; D005399:Fishes / Q000196:embryology / Q000235:genetics*; D016678:Genome; D023281:Genomics; D059014:High-Throughput Nucleotide Sequencing; D023421:Models, Animal; D010802:Phylogeny; D026801:Synteny; D059467:Transcriptome
references.citation: J Exp Zool B Mol Dev Evol. 2014 Sep;322(6):322-33
references.pmid: 23908136
references.citation: Genome Res. 2014 Apr;24(4):688-96
references.pmid: 24418700
references.citation: Nucleic Acids Res. 2013 Jan;41(Database issue):D700-5
references.pmid: 23193262
references.citation: Proc Natl Acad Sci U S A. 2011 Dec 27;108(52):21146-51
references.pmid: 22160688
references.citation: Genome Res. 2004 Jan;14(1):11-7
references.pmid: 14707166
references.citation: Curr Biol. 2007 Mar 6;17(5):452-4
references.pmid: 17306543
references.citation: Gene. 2009 Feb 1;430(1-2):5-11
references.pmid: 19027055
references.citation: Genetica. 2007 Jan;129(1):105-26
references.pmid: 16897450
references.citation: Curr Biol. 2010 Oct 12;20(19):1729-34
references.pmid: 20869245
references.citation: Evodevo. 2013 Jan 23;4(1):3
references.pmid: 23342976
references.citation: J Exp Zool B Mol Dev Evol. 2007 Sep 15;308(5):578-90
references.pmid: 17094081
references.citation: Nature. 2007 Jun 7;447(7145):714-9
references.pmid: 17554307
references.citation: Genome Biol Evol. 2009 Nov 25;1:479-93
references.pmid: 20333216
references.citation: BMC Genomics. 2011 Apr 20;12:202
references.pmid: 21507250
references.citation: BMC Dev Biol. 2001;1:6
references.pmid: 11319037
references.citation: Nat Genet. 2013 May;45(5):567-72
references.pmid: 23542700
references.citation: Nature. 2013 Apr 25;496(7446):494-7
references.pmid: 23594742
references.citation: PLoS One. 2013 Jul 11;8(7):e68389
references.pmid: 23874606
references.citation: Mol Phylogenet Evol. 2003 Jan;26(1):110-20
references.pmid: 12470943
references.citation: BMC Genomics. 2013 Jan 16;14:32
references.pmid: 23324215
references.citation: Nat Commun. 2013;4:1958
references.pmid: 23739623
references.citation: Cold Spring Harb Protoc. 2009 May;2009(5):pdb.emo117
references.pmid: 20147149
references.citation: Comp Biochem Physiol Part D Genomics Proteomics. 2013 Mar;8(1):11-6
references.pmid: 23201534
references.citation: Heredity (Edinb). 2010 Nov;105(5):413-22
references.pmid: 20068586
references.citation: PLoS One. 2013 Oct 25;8(10):e77647
references.pmid: 24204902
references.citation: Nat Genet. 2014 Mar;46(3):253-60
references.pmid: 24487278
references.citation: Evolution. 2009 Jun;63(6):1574-92
references.pmid: 19222565
references.citation: PLoS One. 2013;8(1):e53553
references.pmid: 23326453
references.citation: G3 (Bethesda). 2011 Aug;1(3):171-82
references.pmid: 22384329
references.citation: OMICS. 2014 Feb;18(2):98-110
references.pmid: 24380445
references.citation: Mol Biol Evol. 2006 Jan;23(1):121-36
references.pmid: 16162861
references.citation: J Exp Biol. 2002 Mar;205(Pt 6):769-79
references.pmid: 11914385
references.citation: Proc Biol Sci. 2007 Feb 22;274(1609):489-98
references.pmid: 17476768
references.citation: Nucleic Acids Res. 2013 Jan;41(Database issue):D854-60
references.pmid: 23074187
references.citation: Nucleic Acids Res. 2013 Aug;41(15):e151
references.pmid: 23814184
references.citation: Proc Natl Acad Sci U S A. 2013 Oct 15;110(42):16933-8
references.pmid: 24085851
references.citation: BMC Evol Biol. 2007 Mar 20;7:44
references.pmid: 17374158
references.citation: Nat Commun. 2014 Apr 22;5:3657
references.pmid: 24755649
references.citation: Syst Biol. 2008 Aug;57(4):519-39
references.pmid: 18622808
references.citation: Development. 2014 Jan;141(2):318-24
references.pmid: 24306107
references.citation: Genome Res. 2013 Oct;23(10):1740-8
references.pmid: 23878157
references.citation: Nat Rev Genet. 2007 Mar;8(3):206-16
references.pmid: 17304246
references.citation: J Exp Zool B Mol Dev Evol. 2007 Sep 15;308(5):563-77
references.pmid: 17068775
references.citation: PLoS Biol. 2005 Jan;3(1):e7
references.pmid: 15630479
references.citation: Bioessays. 2008 Sep;30(9):822-32
references.pmid: 18693261
references.citation: Development. 2013 Oct;140(20):4287-95
references.pmid: 24048591
references.citation: Nature. 2010 Jul 8;466(7303):234-7
references.pmid: 20574421
references.citation: BMC Genomics. 2013 Jul 17;14:482
references.pmid: 23865519
references.citation: Mol Ecol. 2011 Nov;20(22):4707-21
references.pmid: 21951675
references.citation: BMC Evol Biol. 2012 Mar 05;12:27
references.pmid: 22390748
references.citation: Mol Ecol. 2013 Feb;22(3):620-34
references.pmid: 22998089
references.citation: Science. 2010 Jan 15;327(5963):302-5
references.pmid: 20007865
references.citation: Mol Ecol. 2009 Jan;18(2):169-85
references.pmid: 18992003
references.citation: Science. 2012 Feb 10;335(6069):677
references.pmid: 22323812
references.citation: Mol Biol Evol. 2005 Dec;22(12):2417-27
references.pmid: 16093565
references.citation: Genome Res. 2003 Mar;13(3):382-90
references.pmid: 12618368
references.citation: PLoS One. 2013;8(3):e59571
references.pmid: 23527220
references.citation: G3 (Bethesda). 2014 Mar 20;4(3):433-45
references.pmid: 24408034
references.citation: Trends Genet. 2007 Mar;23(3):108-12
references.pmid: 17275132
references.citation: J Fish Biol. 2011 Dec;79(6):1545-62
references.pmid: 22136239
references.citation: BMC Evol Biol. 2009 Jun 05;9:127
references.pmid: 19500364
references.citation: Curr Zool. 2013 Apr 1;59(2):170-174
references.pmid: 24363664
references.citation: Mar Genomics. 2012 Sep;7:3-6
references.pmid: 22897955
references.citation: PLoS Curr. 2013 Apr 16;5:
references.pmid: 23788273
references.citation: Trends Ecol Evol. 2005 Jun;20(6):312-9
references.pmid: 16701387
references.citation: Integr Comp Biol. 2010 Dec;50(6):1009-17
references.pmid: 21082069
references.citation: Methods Mol Biol. 2012;888:235-60
references.pmid: 22665285
references.citation: Trends Genet. 2006 Dec;22(12):654-61
references.pmid: 17034900
references.citation: Genome Res. 2009 Feb;19(2):327-35
references.pmid: 19029536
references.citation: Science. 1998 Nov 27;282(5394):1711-4
references.pmid: 9831563
references.citation: Dev Dyn. 2005 Dec;234(4):815-23
references.pmid: 16252286
references.citation: PLoS One. 2012;7(2):e32231
references.pmid: 22384188
references.citation: Gene. 2014 Aug 10;546(2):386-9
references.pmid: 24835514
references.citation: Dev Biol. 2005 Jul 1;283(1):97-112
references.pmid: 15890331
references.citation: Dev Biol. 2009 Aug 1;332(1):90-102
references.pmid: 19433081
references.citation: J Exp Zool B Mol Dev Evol. 2014 Sep;322(6):379-89
references.pmid: 24038780
references.citation: Genome Res. 2012 Mar;22(3):577-91
references.pmid: 22110045
references.citation: Mol Ecol. 2013 Jun;22(11):3036-48
references.pmid: 23181719
references.citation: Proc Natl Acad Sci U S A. 2012 Jan 10;109(2):529-34
references.pmid: 22203992
references.citation: PLoS Genet. 2013 May;9(5):e1003561
references.pmid: 23737760
references.citation: BMC Genomics. 2013 Apr 02;14:214
references.pmid: 23547919
references.citation: Proc Biol Sci. 2006 Apr 22;273(1589):895-9
references.pmid: 16627273
references.citation: Proc Natl Acad Sci U S A. 2013 Aug 20;110(34):13904-9
references.pmid: 23918387
references.citation: BMC Genomics. 2013 Nov 20;14:805
references.pmid: 24252228
references.citation: BMC Biol. 2013 May 09;11:56
references.pmid: 23659605
references.citation: Genetics. 2013 Dec;195(4):1337-52
references.pmid: 24077304
references.citation: Genome Res. 2009 Aug;19(8):1404-18
references.pmid: 19439512
references.citation: BMC Genomics. 2013 Mar 16;14:185
references.pmid: 23496936
references.citation: PLoS One. 2012;7(5):e37880
references.pmid: 22693581
references.citation: Nature. 2011 Aug 10;477(7363):207-10
references.pmid: 21832995
references.citation: Brief Funct Genomics. 2014 Mar;13(2):144-56
references.pmid: 24291769
references.citation: Annu Rev Genet. 2010;44:217-41
references.pmid: 20731603
references.citation: Mech Dev. 2004 Jul;121(7-8):629-37
references.pmid: 15210172
references.citation: Mol Ecol. 2013 Jun;22(11):2986-3001
references.pmid: 23551333
references.citation: Annu Rev Genet. 2009;43:25-47
references.pmid: 19640230
references.citation: Trends Genet. 2004 Oct;20(10):481-90
references.pmid: 15363902
references.citation: Trends Genet. 2003 Dec;19(12):674-8
references.pmid: 14642744
references.citation: Proc Biol Sci. 2009 Jun 22;276(1665):2179-86
references.pmid: 19324738
references.citation: Science. 2011 Aug 19;333(6045):1019-24
references.pmid: 21852499
references.citation: Genetics. 2011 Aug;188(4):799-808
references.pmid: 21828280
references.citation: Mol Biol Evol. 2013 Nov;30(11):2383-400
references.pmid: 23927992
references.citation: Zebrafish. 2013 Mar;10(1):116-8
references.pmid: 23536990
references.citation: Genome Biol. 2010;11(9):403
references.pmid: 20887641
references.citation: FEBS Lett. 2004 Sep 24;575(1-3):119-26
references.pmid: 15388345
references.citation: Zebrafish. 2006;3(1):65-83
references.pmid: 18248247
references.citation: Evol Dev. 2008 May-Jun;10(3):265-72
references.pmid: 18460088
references.citation: Nat Protoc. 2011 Dec 01;6(12):1998-2021
references.pmid: 22134125
references.citation: BMC Genomics. 2013 Sep 21;14:634
references.pmid: 24053439
references.citation: J Morphol. 2013 Jun;274(6):687-703
references.pmid: 23378277
references.citation: Development. 2005 Jan;132(1):89-104
references.pmid: 15563521
references.citation: Nat Rev Genet. 2011 Jun 17;12(7):499-510
references.pmid: 21681211
references.citation: J Appl Ichthyol. 2012 Jun 1;28(3):300-305
references.pmid: 22736877
references.citation: PLoS One. 2013 Oct 10;8(10):e77396
references.pmid: 24130881
references.citation: Mol Ecol. 2012 Jan;21(2):237-49
references.pmid: 21988725
references.citation: Evodevo. 2013 Sep 30;4(1):26
references.pmid: 24079393
references.citation: Philos Trans R Soc Lond B Biol Sci. 2012 Feb 5;367(1587):385-94
references.pmid: 22201168
references.citation: Nucleic Acids Res. 2013 Aug;41(14):e141
references.pmid: 23748566
references.citation: Theory Biosci. 2009 May;128(2):109-20
references.pmid: 19225820
references.citation: Genetics. 2014 Apr;196(4):1303-19
references.pmid: 24496009
references.citation: Pigment Cell Melanoma Res. 2014 Mar;27(2):221-33
references.pmid: 24279354
references.citation: Dis Model Mech. 2014 Feb;7(2):181-92
references.pmid: 24271780
references.citation: PLoS Genet. 2010 Feb 26;6(2):e1000862
references.pmid: 20195501
references.citation: Gen Comp Endocrinol. 2014 Jan 1;195:107-15
references.pmid: 24188887
references.citation: BMC Evol Biol. 2009 Aug 08;9:194
references.pmid: 19664233
references.citation: Trends Ecol Evol. 2000 Jul;15(7):267-271
references.pmid: 10856946
references.citation: Genome Res. 2014 Apr;24(4):604-15
references.pmid: 24487721
references.citation: Respir Physiol. 2000 Aug;122(1):71-80
references.pmid: 10936602
references.citation: J Exp Zool B Mol Dev Evol. 2014 Sep;322(6):403-14
references.pmid: 24106203
references.citation: BMC Genomics. 2013 Nov 11;14:780
references.pmid: 24215161
references.citation: Mol Biol Evol. 2012 Nov;29(11):3529-39
references.pmid: 22734051
references.citation: Science. 2008 Sep 5;321(5894):1346-50
references.pmid: 18772437
references.citation: Gigascience. 2013 Jul 22;2(1):10
references.pmid: 23870653
references.citation: BMC Genomics. 2006 Feb 08;7:20
references.pmid: 16466575
references.citation: Genome Biol Evol. 2012;4(9):937-53
references.pmid: 22851613
references.citation: J Mol Evol. 2004 Aug;59(2):190-203
references.pmid: 15486693
references.citation: Nature. 2004 Oct 21;431(7011):946-57
references.pmid: 15496914
references.citation: Evolution. 2013 Aug;67(8):2166-79
references.pmid: 23888843
references.citation: Nat Protoc. 2006;1(3):1297-305
references.pmid: 17406414
references.citation: G3 (Bethesda). 2011 Dec;1(7):531-8
references.pmid: 22384364
references.citation: Philos Trans R Soc Lond B Biol Sci. 2007 Dec 29;362(1488):2215-32
references.pmid: 17553777
references.citation: PLoS Genet. 2009 Jan;5(1):e1000326
references.pmid: 19119422
references.citation: Mol Biol Evol. 2011 Mar;28(3):1205-15
references.pmid: 21081479
references.citation: Science. 2012 Nov 2;338(6107):619-21
references.pmid: 23118176
references.citation: Biol Lett. 2009 Dec 23;5(6):812-6
references.pmid: 19586967
references.citation: Proc Natl Acad Sci U S A. 2012 Feb 28;109(9):3434-9
references.pmid: 22331888
references.citation: Dev Growth Differ. 2008 Aug;50(6):415-9
references.pmid: 18422685
references.citation: Proc Natl Acad Sci U S A. 1997 Apr 1;94(7):3420-4
references.pmid: 9096409
references.citation: Dev Dyn. 2013 Aug;242(8):949-963
references.pmid: 23723152
references.citation: PLoS One. 2012;7(8):e43762
references.pmid: 22916302
references.citation: G3 (Bethesda). 2013 Jan;3(1):65-74
references.pmid: 23316439
references.citation: J Exp Biol. 2004 Oct;207(Pt 21):3649-56
references.pmid: 15371473
references.citation: Genome Res. 2009 Aug;19(8):1497-505
references.pmid: 19465509
references.citation: Genes Nutr. 2013 Jan;8(1):5-17
references.pmid: 22903900
references.citation: J Invest Dermatol. 2010 Jan;130(1):249-58
references.pmid: 19609310
references.citation: PLoS One. 2013;8(2):e55659
references.pmid: 23405189
references.citation: Genetics. 2009 Dec;183(4):1385-95
references.pmid: 19786620
references.citation: Science. 2002 Aug 23;297(5585):1301-10
references.pmid: 12142439
references.citation: J Exp Biol. 2000 Aug;203(Pt 15):2331-9
references.pmid: 10887071
references.citation: PLoS One. 2013 Nov 25;8(11):e80823
references.pmid: 24282555
references.citation: Gene. 2012 Dec 15;511(2):195-201
references.pmid: 23026207
references.citation: J Fish Biol. 2011 Jun;78(6):1624-46
references.pmid: 21651520
references.citation: Curr Opin Genet Dev. 2008 Dec;18(6):544-50
references.pmid: 19095434
references.citation: J Exp Zool B Mol Dev Evol. 2014 Sep;322(6):415-37
references.pmid: 24482296
references.citation: PLoS One. 2008;3(10):e3376
references.pmid: 18852878
references.citation: Proc Natl Acad Sci U S A. 2012 Mar 6;109(10):3838-43
references.pmid: 22355122
references.citation: Nat Commun. 2013;4:2769
references.pmid: 24225650
references.citation: Nature. 1999 Sep 30;401(6752):443-4
references.pmid: 10519545
references.citation: BMC Genomics. 2012 Mar 19;13:96
references.pmid: 22424280
references.citation: Genome Biol. 2007;8 Suppl 1:S10
references.pmid: 18047687
references.citation: Genome Biol Evol. 2013;5(1):45-60
references.pmid: 23196969
references.citation: Nat Rev Genet. 2011 Sep 07;12(10):671-82
references.pmid: 21897427
references.citation: Proc Biol Sci. 2009 Feb 22;276(1657):623-31
references.pmid: 19004755
references.citation: Nat Genet. 2006 Jan;38(1):107-11
references.pmid: 16341223
references.citation: Nat Rev Genet. 2001 May;2(5):333-41
references.pmid: 11331899
references.citation: Proc Natl Acad Sci U S A. 2012 Aug 21;109(34):13698-703
references.pmid: 22869754
references.citation: Nature. 2007 May 24;447(7143):473-6
references.pmid: 17522683
references.citation: Cell. 2013 Feb 14;152(4):691-702
references.pmid: 23415220
references.citation: Development. 2014 Feb;141(3):715-24
references.pmid: 24449846
references.citation: Genetics. 2014 Jun;197(2):625-41
references.pmid: 24700104
references.citation: Comp Biochem Physiol C Toxicol Pharmacol. 2014 Jun;163:24-36
references.pmid: 24486528
references.citation: BMC Genomics. 2010 Nov 16;11:635
references.pmid: 21080939
references.citation: J Exp Zool B Mol Dev Evol. 2011 Sep 15;316(6):451-64
references.pmid: 21688387
references.citation: Nat Rev Genet. 2004 Apr;5(4):288-98
references.pmid: 15131652
references.citation: Nature. 2004 Oct 14;431(7010):844-7
references.pmid: 15483612
references.citation: BMC Genomics. 2013 Aug 20;14:566
references.pmid: 23957753
references.citation: J Exp Zool B Mol Dev Evol. 2014 Sep;322(6):438-63
references.pmid: 24464682
references.citation: Mol Ecol Resour. 2014 May;14(3):622-35
references.pmid: 24354690
references.citation: Zebrafish. 2012 Jun;9(2):59-67
references.pmid: 22715948
references.citation: Methods Cell Biol. 2011;104:259-85
references.pmid: 21924168
references.citation: Methods. 2013 Aug 15;62(3):207-15
references.pmid: 23624103
references.citation: Genetics. 2013 Mar;193(3):739-49
references.pmid: 23288935
references.citation: Genome Res. 2012 Nov;22(11):2241-9
references.pmid: 22800726
references.citation: Mol Biol Evol. 2006 Sep;23(9):1808-16
references.pmid: 16809621
references.citation: Proc Natl Acad Sci U S A. 2008 Sep 2;105(35):12944-9
references.pmid: 18753634
references.citation: Nature. 2012 Apr 04;484(7392):55-61
references.pmid: 22481358
references.citation: J Exp Zool B Mol Dev Evol. 2011 Nov 15;316(7):526-46
references.pmid: 21809437
references.citation: Genome Res. 2004 Jan;14(1):1-10
references.pmid: 14707165
references.citation: BMC Biol. 2012 Dec 27;10:108
references.pmid: 23270452
references.citation: Science. 2013 Dec 13;342(6164):1372-5
references.pmid: 24337296
references.citation: Dev Biol. 2011 Sep 15;357(2):450-62
references.pmid: 21435340
references.citation: Nature. 2013 Apr 25;496(7446):498-503
references.pmid: 23594743
references.citation: Nature. 2013 Apr 18;496(7445):311-6
references.pmid: 23598338
references.citation: Genetics. 2001 Jul;158(3):1203-15
references.pmid: 11454768
references.citation: Genome Res. 2007 Feb;17(2):240-8
references.pmid: 17189378
references.citation: Semin Cell Dev Biol. 2013 Feb;24(2):119-27
references.pmid: 23291292
references.citation: J Exp Zool B Mol Dev Evol. 2007 Dec 15;308(6):757-68
references.pmid: 17849442
references.citation: Nat Rev Genet. 2009 Oct;10(10):725-32
references.pmid: 19652647
references.citation: Dev Dyn. 2011 Sep;240(9):2101-7
references.pmid: 21805532
references.citation: PLoS Curr. 2013 Apr 18;5:
references.pmid: 23653398
references.citation: Nat Methods. 2009 Dec;6(12):911-6
references.pmid: 19898487
references.citation: BMC Evol Biol. 2007 Mar 13;7:38
references.pmid: 17355618
references.citation: Nature. 2011 Jan 6;469(7328):84-8
references.pmid: 21209663
references.citation: PLoS Biol. 2014 Jan;12(1):e1001773
references.pmid: 24465181
references.citation: Nature. 1954 May 8;173(4410):848-50
references.pmid: 13165664
references.citation: Bioessays. 2007 Sep;29(9):884-96
references.pmid: 17691105
references.citation: Cell Res. 2013 Apr;23(4):465-72
references.pmid: 23528705
references.citation: PLoS One. 2013;8(2):e57281
references.pmid: 23437360
references.citation: Mol Biol Evol. 2013 Jul;30(7):1527-43
references.pmid: 23612715
references.citation: Cell. 2008 Jul 11;134(1):25-36
references.pmid: 18614008
references.citation: Genetics. 2013 Jul;194(3):631-46
references.pmid: 23666934
references.citation: Dev Dyn. 2009 Sep;238(9):2409-17
references.pmid: 19653328
references.citation: Trends Genet. 2009 Feb;25(2):74-81
references.pmid: 19108930
references.citation: J Exp Zool B Mol Dev Evol. 2010 Jul 15;314(5):369-81
references.pmid: 20235119
references.citation: Dev Genes Evol. 2001 Jul;211(7):319-28
references.pmid: 11466528
references.citation: Dev Biol. 2009 Jun 1;330(1):200-11
references.pmid: 19285488
references.citation: PLoS One. 2013 Apr 23;8(4):e61378
references.pmid: 23626677
references.citation: Bioinformatics. 2012 Mar 15;28(6):900-4
references.pmid: 22332236