title: Towards an integrated experimental-theoretical approach for assessing the mechanistic basis of hair and feather morphogenesis.
abstract: In his seminal 1952 paper, 'The Chemical Basis of Morphogenesis', Alan Turing lays down a milestone in the application of theoretical approaches to understand complex biological processes. His deceptively simple demonstration that a system of reacting and diffusing chemicals could, under certain conditions, generate spatial patterning out of homogeneity provided an elegant solution to the problem of how one of nature's most intricate events occurs: the emergence of structure and form in the developing embryo. The molecular revolution that has taken place during the six decades following this landmark publication has now placed this generation of theoreticians and biologists in an excellent position to rigorously test the theory and, encouragingly, a number of systems have emerged that appear to conform to some of Turing's fundamental ideas. In this paper, we describe the history and more recent integration between experiment and theory in one of the key models for understanding pattern formation: the emergence of feathers and hair in the skins of birds and mammals.
authors.affiliation: Department of Mathematics/Maxwell Institute for Mathematical Sciences , Heriot-Watt University , Edinburgh EH14 4AS , UK.
authors.forename: K J
authors.initials: KJ
authors.lastname: Painter
authors.forename: G S
authors.initials: GS
authors.lastname: Hunt
authors.forename: K L
authors.initials: KL
authors.lastname: Wells
authors.forename: J A
authors.initials: JA
authors.lastname: Johansson
authors.forename: D J
authors.initials: DJ
authors.lastname: Headon
keywords: Turing patterns; activator–inhibitor; feather buds and hair follicles; morphogenesis; reaction–diffusion; skin patterning
references.citation: J Theor Biol. 1985 May 21;114(2):243-72
references.pmid: 4033155
references.citation: J Math Biol. 2009 Jan;58(1-2):183-217
references.pmid: 18626644
references.citation: Dev Biol. 2009 Oct 15;334(2):369-82
references.pmid: 19647731
references.citation: Dev Cell. 2009 Jul;17(1):49-61
references.pmid: 19619491
references.citation: Kybernetik. 1972 Dec;12(1):30-9
references.pmid: 4663624
references.citation: Trends Cell Biol. 2009 Oct;19(10):523-30
references.pmid: 19733079
references.citation: Nature. 1995 Aug 31;376(6543):765-8
references.pmid: 24547605
references.citation: Int J Dev Biol. 2004;48(2-3):117-35
references.pmid: 15272377
references.citation: Development. 2007 Mar;134(5):1011-22
references.pmid: 17267445
references.citation: Philos Trans R Soc Lond B Biol Sci. 1981 Oct 7;295(1078):473-96
references.pmid: 6117906
references.citation: J Theor Biol. 1972 Dec;37(3):545-59
references.pmid: 4645361
references.citation: Bull Math Biol. 2012 Apr;74(4):935-57
references.pmid: 22072186
references.citation: Bull Math Biol. 1999 Nov;61(6):1093-120
references.pmid: 17879872
references.citation: Science. 1999 Jun 11;284(5421):1826-8
references.pmid: 10364553
references.citation: Nature. 1991 Feb 14;349(6310):630-3
references.pmid: 2000137
references.citation: Dev Cell. 2006 Oct;11(4):495-504
references.pmid: 17011489
references.citation: Bioessays. 2000 Aug;22(8):753-60
references.pmid: 10918306
references.citation: Nat Rev Mol Cell Biol. 2011 Aug 18;12(9):594-604
references.pmid: 21850035
references.citation: J Theor Biol. 2006 Jun 21;240(4):562-73
references.pmid: 16364368
references.citation: Dev Biol. 1998 Apr 1;196(1):11-23
references.pmid: 9527877
references.citation: J Math Biol. 2011 Dec;63(6):1095-119
references.pmid: 21274536
references.citation: J Theor Biol. 1979 Dec 7;81(3):389-400
references.pmid: 537379
references.citation: Int J Dev Biol. 2009;53(5-6):783-94
references.pmid: 19557684
references.citation: J Theor Biol. 2006 Nov 7;243(1):98-113
references.pmid: 16860344
references.citation: Dev Biol. 2000 Jan 1;217(1):54-61
references.pmid: 10625535
references.citation: J Theor Biol. 1969 Oct;25(1):1-47
references.pmid: 4390734
references.citation: Bull Math Biol. 2010 Nov;72(8):2139-60
references.pmid: 20309645
references.citation: Bioessays. 2005 Mar;27(3):247-61
references.pmid: 15714560
references.citation: C R Biol. 2003 Aug;326(8):717-27
references.pmid: 14608692
references.citation: Mech Dev. 1999 Aug;86(1-2):51-62
references.pmid: 10446265
references.citation: J Math Biol. 1983;17(1):125-9
references.pmid: 6875405
references.citation: Dev Biol. 2007 May 15;305(2):498-507
references.pmid: 17397822
references.citation: Exp Dermatol. 2006 Dec;15(12):931-49
references.pmid: 17083360
references.citation: Biophys J. 1995 May;68(5):2181-9
references.pmid: 7612862
references.citation: J Math Biol. 2010 Jul;61(1):133-64
references.pmid: 19727733
references.citation: Nature. 1980 Oct 30;287(5785):795-801
references.pmid: 6776413
references.citation: J Cell Sci. 2009 Sep 15;122(Pt 18):3215-23
references.pmid: 19726631
references.citation: Dev Biol. 2008 Aug 1;320(1):60-71
references.pmid: 18508042
references.citation: C R Biol. 2003 Feb;326(2):239-52
references.pmid: 12754942
references.citation: Proc Natl Acad Sci U S A. 2006 Jun 13;103(24):9075-80
references.pmid: 16769906
references.citation: J Theor Biol. 1982 Oct 21;98(4):575-607
references.pmid: 7154690
references.citation: Science. 2007 Jan 26;315(5811):521-5
references.pmid: 17255514
references.citation: Trends Cogn Sci. 2009 Jan;13(1):36-43
references.pmid: 19058992
references.citation: Science. 2008 Oct 17;322(5900):399-403
references.pmid: 18927385
references.citation: Mech Dev. 1999 Dec;89(1-2):87-92
references.pmid: 10559483
references.citation: IMA J Math Appl Med Biol. 1992;9(4):227-48
references.pmid: 1302759
references.citation: J Embryol Exp Morphol. 1983 Dec;78:83-125
references.pmid: 6663234
references.citation: J Cell Sci. 1974 Jul;15(2):321-46
references.pmid: 4859215
references.citation: J Theor Biol. 1970 Mar;26(3):399-415
references.pmid: 5462335
references.citation: Phys Rev E Stat Nonlin Soft Matter Phys. 2009 Mar;79(3 Pt 1):031908
references.pmid: 19391972
references.citation: Phys Rev Lett. 1990 Jun 11;64(24):2953-2956
references.pmid: 10041855
references.citation: Nature. 1989 Sep 28;341(6240):282-3
references.pmid: 2797143
references.citation: J Theor Biol. 1996 Dec 21;183(4):429-46
references.pmid: 9015458
references.citation: Dev Cell. 2002 Sep;3(3):425-37
references.pmid: 12361604
references.citation: Science. 2006 Dec 1;314(5804):1447-50
references.pmid: 17082421
references.citation: Proc Natl Acad Sci U S A. 1999 May 11;96(10):5549-54
references.pmid: 10318921
references.citation: Bull Math Biol. 2006 Nov;68(8):1975-2003
references.pmid: 17086491
references.citation: J Theor Biol. 1971 Sep;32(3):507-37
references.pmid: 5571122
references.citation: Development. 2010 Jul;137(14):2253-64
references.pmid: 20570935
references.citation: J Theor Biol. 1964 Sep;7:302-17
references.pmid: 14212276
references.citation: Science. 2010 Sep 24;329(5999):1616-20
references.pmid: 20929839
references.citation: Proc Biol Sci. 2002 Apr 22;269(1493):781-92
references.pmid: 11958709
references.citation: PLoS Biol. 2011 Mar;9(3):e1001028
references.pmid: 21423653
references.citation: Nature. 1995 Jul 6;376(6535):49-53
references.pmid: 7596432
references.citation: Development. 2011 May;138(9):1807-16
references.pmid: 21447550
references.citation: Phys Rev E Stat Phys Plasmas Fluids Relat Interdiscip Topics. 1993 Mar;47(3):2128-2154
references.pmid: 9960234
references.citation: Bull Math Biol. 2006 Jan;68(1):99-130
references.pmid: 16794923
references.citation: Biophys J. 2010 Aug 9;99(4):L33-5
references.pmid: 20712981
references.citation: Bull Math Biol. 2002 Jul;64(4):747-69
references.pmid: 12216419
references.citation: Science. 1991 Feb 8;251(4994):650-2
references.pmid: 17741380
references.citation: Phys Rev E Stat Nonlin Soft Matter Phys. 2007 Jul;76(1 Pt 1):011902
references.pmid: 17677489
references.citation: Math Model Nat Phenom. 2009 Jan 1;4(4):3-82
references.pmid: 19844610
references.citation: J Theor Biol. 1985 Jul 21;115(2):299-317
references.pmid: 2412073
references.citation: Proc Natl Acad Sci U S A. 2005 Aug 16;102(33):11734-9
references.pmid: 16087884
references.citation: J Math Biol. 1988;26(1):27-39
references.pmid: 3351393
references.citation: Development. 2008 Aug;135(16):2797-805
references.pmid: 18635609
references.citation: Int J Dev Biol. 2004;48(2-3):181-91
references.pmid: 15272383
references.citation: Bull Math Biol. 1999 May;61(3):483-505
references.pmid: 17883228
references.citation: Science. 1978 Jan 20;199(4326):259-70
references.pmid: 413193
references.citation: Science. 2009 May 8;324(5928):772-5
references.pmid: 19423823
references.citation: Mol Biosyst. 2009 Dec;5(12):1549-68
references.pmid: 20023719
references.citation: Curr Opin Genet Dev. 2007 Aug;17(4):281-6
references.pmid: 17624758
references.citation: J Theor Biol. 1971 Feb;30(2):225-34
references.pmid: 4926701
references.citation: Development. 1999 Nov;126(22):4997-5009
references.pmid: 10529418
references.citation: Annu Rev Microbiol. 1998;52:779-806
references.pmid: 9891813
references.citation: Nature. 2011 May 19;473(7347):337-42
references.pmid: 21593866
references.citation: J Invest Dermatol. 2002 Feb;118(2):216-25
references.pmid: 11841536
references.citation: Development. 2011 Jan;138(2):273-82
references.pmid: 21148187
references.citation: J Math Biol. 1986;24(2):141-65
references.pmid: 3755746
references.citation: Cell. 2007 Jul 13;130(1):141-52
references.pmid: 17632061
references.citation: J Theor Biol. 1973 Aug 15;40(2):353-67
references.pmid: 4747247
references.citation: J Embryol Exp Morphol. 1961 Dec;9:661-72
references.pmid: 13914305
references.citation: Trends Genet. 1996 Sep;12(9):359-64
references.pmid: 8855666
references.citation: Dev Dyn. 1992 Nov;195(3):201-15
references.pmid: 1301084