The human crystallin gene families

Human Genomics

Volumn 6, Issue 1, 2012, Pages

  Wistow, Graeme ^a  

^a NATIONAL EYE INSTITUTE   (United States)

Author keywords

Cataract; Crystallins; Enzymes; Evolution; Heat shock; Lens; Pseudogene

Indexed keywords

ALPHA CRYSTALLIN; BETA CRYSTALLIN; BETAINE HOMOCYSTEINE METHYLTRANSFERASE; CRYSTALLIN; CRYAA PROTEIN, HUMAN; CRYAB PROTEIN, HUMAN; SMALL HEAT SHOCK PROTEIN;

AMINO ACID SEQUENCE; AMINO ACID SUBSTITUTION; ARTICLE; CATARACTOGENESIS; CRYBA1 GENE; CRYBA2 GENE; CRYBA4 GENE; CRYBB1 GENE; CRYBB2 GENE; CRYBB2P1 GENE; CRYBB3 GENE; CRYGA GENE; CRYGFP GENE; CRYGS GENE; CRYSTALLIN GENE; CYRGGP GENE; EXON; EXPRESSED SEQUENCE TAG; GENE; GENE CLUSTER; GENE EXPRESSION; GENE MUTATION; GENE STRUCTURE; HUMAN; LENS; NONHUMAN; NONSENSE MUTATION; OPEN READING FRAME; PHYLOGENETIC TREE; PROTEIN MODIFICATION; CATARACT; GENETICS; METABOLISM; MULTIGENE FAMILY; MUTATION; PATHOPHYSIOLOGY; REVIEW;

ALPHA-CRYSTALLIN B CHAIN; BETAINE-HOMOCYSTEINE S-METHYLTRANSFERASE; CATARACT; CRYSTALLINS; HEAT-SHOCK PROTEINS, SMALL; HUMANS; LENS, CRYSTALLINE; MULTIGENE FAMILY; MUTATION;

MLCS; MLOWN;

EID: 84876513077     PISSN: 14739542     EISSN: 14797364     Source Type: Journal    
DOI: 10.1186/1479-7364-6-26     Document Type: Article

References (95)

1. Wistow G, Slingsby C: Structure and evolution of crystallins. The Encyclopedia of the Eye 2010, 2:229-238.
2. Bloemendal H, De Jong W, Jaenicke R, Lubsen NH, Slingsby C, Tardieu A: Ageing and vision: structure, stability and function of lens crystallins. Prog Biophys Mol Biol 2004, 86:407-485.
3. Harding JJ, Crabbe MJC: The lens: development, proteins, metabolism and cataract. Eye 1984, 1B:207-492.
4. Wistow G: Lens crystallins: gene recruitment and evolutionary dynamism. Trends Biochem Sci 1993, 18:301-306.
5. Piatigorsky J: Review: a case for corneal crystallins. J Ocul Pharmacol Ther 2000, 16:173-180.
6. Lassen N, Black WJ, Estey T, Vasiliou V: The role of corneal crystallins in the cellular defense mechanisms against oxidative stress. Semin Cell Dev Biol 2008, 19:100-112.
7. Jester JV: Corneal crystallins and the development of cellular transparency. Semin Cell Dev Biol 2008, 19:82-93.
8. Wistow G, Peterson K, Gao J, Buchoff P, Jaworski C, Bowes-Rickman C, Ebright JN, Hauser MA, Hoover D: NEIBank: genomics and bioinformatics resources for vision research. Mol Vis 2008, 14:1327-1337.
9. Clark AR, Lubsen NH, Slingsby C: sHSP in the eye lens: crystallin mutations, cataract and proteostasis. Int J Biochem Cell Biol 2012, 44:1687-1697.
10. van Dijk MA, Sweers MA, de Jong WW: The evolution of an alternatively spliced exon in the alphaA-crystallin gene. J Mol Evol 2001, 52:510-515.
11. Jaworski CJ, Piatigorsky J: A pseudo-exon in the functional human alphaA-crystallin gene. Nature 1989, 337:752-754.
12. de Jong WW, Caspers GJ, Leunissen JA: Genealogy of the alphacrystallin-small heat-shock protein superfamily. Int J Biol Macromol 1998, 22:151-162.
13. Fontaine JM, Rest JS, Welsh MJ, Benndorf R: The sperm outer dense fiber protein is the 10th member of the superfamily of mammalian small stress proteins. Cell Stress Chaperones 2003, 8:62-69.
14. Kappe G, Boelens WC, de Jong WW: Why proteins without an alphacrystallin domain should not be included in the human small heat shock protein family HSPB. Cell Stress Chaperones 2010, 15:457-461.
15. Tamura K, Dudley J, Nei M, Kumar S: MEGA4: Molecular Evolutionary Genetics Analysis (MEGA) software version 4.0. Mol Biol Evol 2007, 24:1596-1599.
16. Rao NA, Saraswathy S, Pararajasegaram G, Bhat SP: Small heat shock protein alphaA-crystallin prevents photoreceptor degeneration in experimental autoimmune uveitis. PLoS One 2012, 7:e33582.
17. Quinlan R: Cytoskeletal competence requires protein chaperones. Prog Mol Subcell Biol 2002, 28:219-233.
18. Mackay DS, Andley UP, Shiels A: Cell death triggered by a novel mutation in the alphaA-crystallin gene underlies autosomal dominant cataract linked to chromosome 21q. Eur J Hum Genet 2003, 11:784-793.
19. Litt M, Kramer P, LaMorticella DM, Murphey W, Lovrien EW, Weleber RG: Autosomal dominant congenital cataract associated with a missense mutation in the human alpha crystallin gene CRYAA. Hum Mol Genet 1998, 7:471-474.
20. Richter L, Flodman P, Barria von-Bischhoffshausen F, Burch D, Brown S, Nguyen L, Turner J, Spence MA, Bateman JB: Clinical variability of autosomal dominant cataract, microcornea and corneal opacity and novel mutation in the alpha A crystallin gene (CRYAA). Am J Med Genet A 2008, 146:833-842.
21. Pras E, Frydman M, Levy-Nissenbaum E, Bakhan T, Raz J, Assia EI, Goldman B: A nonsense mutation (W9X) in CRYAA causes autosomal recessive cataract in an inbred Jewish Persian family. Invest Ophthalmol Vis Sci 2000, 41:3511-3515.
22. Sun W, Xiao X, Li S, Guo X, Zhang Q: Mutation analysis of 12 genes in Chinese families with congenital cataracts. Mol Vis 2011, 17:2197-2206.
23. Selcen D, Engel AG: Myofibrillar myopathy caused by novel dominant negative alpha B-crystallin mutations. Ann Neurol 2003, 54:804-810.
24. Del Bigio MR, Chudley AE, Sarnat HB, Campbell C, Goobie S, Chodirker BN, Selcen D: Infantile muscular dystrophy in Canadian aboriginals is an alphaB-crystallinopathy. Ann Neurol 2011, 69:866-871.
25. Vicart P, Caron A, Guicheney P, Li Z, Prevost MC, Faure A, Chateau D, Chapon F, Tome F, Dupret JM, Paulin D, Fardeau M: A missense mutation in the alphaB-crystallin chaperone gene causes a desmin-related myopathy. Nat Genet 1998, 20:92-95.
26. Berry V, Francis P, Reddy MA, Collyer D, Vithana E, MacKay I, Dawson G, Carey AH, Moore A, Bhattacharya SS, Quinlan RA: Alpha-B crystallin gene (CRYAB) mutation causes dominant congenital posterior polar cataract in humans. Am J Hum Genet 2001, 69:1141-1145.
27. Wistow GJ, Piatigorsky J: Lens crystallins: the evolution and expression of proteins for a highly specialized tissue. Annu Rev Biochem 1988, 57:479-504.
28. Wistow G, Wyatt K, David L, Gao C, Bateman O, Bernstein S, Tomarev S, Segovia L, Slingsby C, Vihtelic T: gammaN-crystallin and the evolution of the betagamma-crystallin superfamily in vertebrates. FEBS J 2005, 272:2276-2291.
29. Hogg D, Tsui LC, Gorin M, Breitman ML: Characterization of the human beta-crystallin gene Hu beta A3/A1 reveals ancestral relationships among the beta gamma-crystallin superfamily. J Biol Chem 1986, 261:12420-12427.
30. Kannabiran C, Rogan PK, Olmos L, Basti S, Rao GN, Kaiser-Kupfer M, Hejtmancik JF: Autosomal dominant zonular cataract with sutural opacities is associated with a splice mutation in the betaA3/A1-crystallin gene. Mol Vis 1998, 4:21.
31. Bateman JB, Geyer DD, Flodman P, Johannes M, Sikela J, Walter N, Moreira AT, Clancy K, Spence MA: A new betaA1-crystallin splice junction mutation in autosomal dominant cataract. Invest Ophthalmol Vis Sci 2000, 41:3278-3285.
32. Yang Z, Li Q, Ma Z, Guo Y, Zhu S, Ma X: A G->T splice site mutation of CRYBA1/A3 associated with autosomal dominant suture cataracts in a Chinese family. Mol Vis 2011, 17:2065-2071.
33. Yang Z, Su D, Li Q, Yang F, Ma Z, Zhu S, Ma X: A novel T->G splice site mutation of CRYBA1/A3 associated with autosomal dominant nuclear cataracts in a Chinese family. Mol Vis 2012, 18:1283-1288.
34. Qi Y, Jia H, Huang S, Lin H, Gu J, Su H, Zhang T, Gao Y, Qu L, Li D, Li Y: A deletion mutation in the betaA1/A3 crystallin gene (CRYBA1/A3) is associated with autosomal dominant congenital nuclear cataract in a Chinese family. Hum Genet 2004, 114:192-197.
35. Sinha D, Valapala M, Bhutto I, Patek B, Zhang C, Hose S, Yang F, Cano M, Stark WJ, Lutty GA, Zigler JS, Wawrousek EF: betaA3/A1-crystallin is required for proper astrocyte template formation and vascular remodeling in the retina. Transgenic Res 2012, 21:1033-1042.
36. Wistow G, Bernstein SL, Wyatt MK, Behal A, Touchman JW, Bouffard G, Smith D, Peterson K: Expressed sequence tag analysis of adult human lens for the NEIBank Project: over 2000 non-redundant transcripts, novel genes and splice variants. Mol Vis 2002, 8:171-184.
37. Lapko VN, Smith DL, Smith JB: Expression of betaA2-crystallin in human lenses. Exp Eye Res 2003, 77:383-385.
38. Duncan MK, Haynes JI, Piatigorsky J: The chicken betaA4- and betaB1-crystallin-encoding genes are tightly linked. Gene 1995, 62:189-196.
39. Billingsley G, Santhiya ST, Paterson AD, Ogata K, Wodak S, Hosseini SM, Manisastry SM, Vijayalakshmi P, Gopinath PM, Graw J, Heon E: CRYBA4, a novel human cataract gene is also involved in microphthalmia. Am J Hum Genet 2006, 79:702-709.
40. Zhou G, Zhou N, Hu S, Zhao L, Zhang C, Qi Y: A missense mutation in CRYBA4 associated with congenital cataract and microcornea. Mol Vis 2010, 16:1019-1024.
41. Mackay DS, Boskovska OB, Knopf HL, Lampi KJ, Shiels A: A nonsense mutation in CRYBB1 associated with autosomal dominant cataract linked to human chromosome 22q. Am J Hum Genet 2002, 71:1216-1221.
42. Cohen D, Bar-Yosef U, Levy J, Gradstein L, Belfair N, Ofir R, Joshua S, Lifshitz T, Carmi R, Birk OS: Homozygous CRYBB1 deletion mutation underlies autosomal recessive congenital cataract. Invest Ophthalmol Vis Sci 2007, 48:2208-2213.
43. Yang J, Zhu Y, Gu F, He X, Cao Z, Li X, Tong Y, Ma X: A novel nonsense mutation in CRYBB1 associated with autosomal dominant congenital cataract. Mol Vis 2008, 14:727-731.
44. Wang KJ, Wang BB, Zhang F, Zhao Y, Ma X, Zhu SQ: Novel beta-crystallin gene mutations in Chinese families with nuclear cataracts. Arch Ophthalmol 2011, 129:337-343.
45. Wang J, Ma X, Gu F, Liu NP, Hao XL, Wang KJ, Wang NL, Zhu SQ: A missense mutation S228P in the CRYBB1 gene causes autosomal dominant congenital cataract. Chin Med J (Engl) 2007, 120:820-824.
46. Willoughby CE, Shafiq A, Ferrini W, Chan LL, Billingsley G, Priston M, Mok C, Chandna A, Kaye S, Heon E: CRYBB1 mutation associated with congenital cataract and microcornea. Mol Vis 2005, 11:587-593.
47. Wang KJ, Wang S, Cao NQ, Yan YB, Zhu SQ: A novel mutation in CRYBB1 associated with congenital cataract-microcornea syndrome: the p. Ser129Arg mutation destabilizes the betaB1/betaA3-crystallin heteromer but not the betaB1-crystallin homomer. Hum Mutat 2011, 32:E2050-E2060.
48. Litt M, Carrero-Valenzuela R, LaMorticella DM, Schultz DW, Mitchell TN, Kramer P, Maumenee IH: Autosomal dominant cerulean cataract is associated with a chain termination mutation in the human betacrystallin gene CRYBB2. Hum Mol Genet 1997, 6:665-668.
49. Gill D, Klose R, Munier FL, McFadden M, Priston M, Billingsley G, Ducrey N, Schorderet DF, Heon E: Genetic heterogeneity of the Coppock-like cataract: a mutation in CRYBB2 on chromosome 22q11.2. Invest Ophthalmol Vis Sci 2000, 41:159-165.
50. Vanita, Sarhadi V, Reis A, Jung M, Singh D, Sperling K, Singh JR, Burger J: A unique form of autosomal dominant cataract explained by gene conversion between beta-crystallin B2 and its pseudogene. J Med Genet 2001, 38:392-396.
51. Bateman JB, von-Bischhoffshaunsen FR, Richter L, Flodman P, Burch D, Spence M: Gene conversion mutation in crystallin, beta-B2 (CRYBB2) in a Chilean family with autosomal dominant cataract. Ophthalmology 2007, 114:425-432.
52. Kumar KD, Kumar GS, Santhiya ST: Nonspecific PCR amplification of CRYBB2-pseudogene leads to misconception of natural variation as mutation. Investig Ophthalmol Vis Sci 2012, 53:5770.
53. Hansen L, Mikkelsen A, Nurnberg P, Nurnberg G, Anjum I, Eiberg H, Rosenberg T: Comprehensive mutational screening in a cohort of Danish families with hereditary congenital cataract. Invest Ophthalmol Vis Sci 2009, 50:3291-3303.
54. Yao K, Li J, Jin C, Wang W, Zhu Y, Shentu X, Wang Q: Characterization of a novel mutation in the CRYBB2 gene associated with autosomal dominant congenital posterior subcapsular cataract in a Chinese family. Mol Vis 2011, 17:144-152.
55. Weisschuh N, Aisenbrey S, Wissinger B, Riess A: Identification of a novel CRYBB2 missense mutation causing congenital autosomal dominant cataract. Mol Vis 2012, 18:174-180.
56. Faletra F, d'Adamo AP, Pensiero S, Athanasakis E, Catalano D, Bruno I, Gasparini P: A novel CRYBB2 missense mutation causing Congenital autosomal dominant cataract in an Italian family. Ophthalmic Genet, in press.
57. Riazuddin SA, Yasmeen A, Yao W, Sergeev YV, Zhang Q, Zulfiqar F, Riaz A, Riazuddin S, Hejtmancik JF: Mutations in betaB3-crystallin associated with autosomal recessive cataract in two Pakistani families. Invest Ophthalmol Vis Sci 2005, 46:2100-2106.
58. Heon E, Priston M, Schorderet DF, Billingsley GD, Girard PO, Lubsen N, Munier FL: The gamma-crystallins and human cataracts: a puzzle made clearer. Am J Hum Genet 1999, 65:1261-1267.
59. Brakenhoff RH, Aarts HJ, Reek FH, Lubsen NH, Schoenmakers JG: Human gamma-crystallin genes. A gene family on its way to extinction. J Mol Biol 1990, 216:519-532.
60. Simpanya MF, Wistow G, Gao J, David LL, Giblin FJ, Mitton KP: Expressed sequence tag analysis of guinea pig (Cavia porcellus) eye tissues for NEIBank. Mol Vis 2008, 14:2413-2427.
61. Brakenhoff RH, Henskens HAM, van Rossum MWPC, Lubsen NH, Schoenmakers JGG: Activation of the gammaE-crystallin pseudogene in the human hereditary Coppock-like cataract. Hum Mol Genet 1994, 3:279-283.
62. Robinson NE, Lampi KJ, Speir JP, Kruppa G, Easterling M, Robinson AB: Quantitative measurement of young human eye lens crystallins by direct injection Fourier transform ion cyclotron resonance mass spectrometry. Mol Vis 2006, 12:704-711.
63. Lapko VN, Smith DL, Smith JB: Methylation and carbamylation of human gamma-crystallins. Protein Sci 2003, 12:1762-1774.
64. Santhiya ST, Shyam Manohar M, Rawlley D, Vijayalakshmi P, Namperumalsamy P, Gopinath PM, Loster J, Graw J: Novel mutations in the gamma-crystallin genes cause autosomal dominant congenital cataracts. J Med Genet 2002, 39:352-358.
65. Ren Z, Li A, Shastry BS, Padma T, Ayyagari R, Scott MH, Parks MM, Kaiser-Kupfer MI, Hejtmancik JF: A 5-base insertion in the gammaCcrystallin gene is associated with autosomal dominant variable zonular pulverulent cataract. Hum Genet 2000, 106:531-537.
66. Stephan DA, Gillanders E, Vanderveen D, Freas-Lutz D, Wistow G, Baxevanis AD, Robbins CM, VanAuken A, Quesenberry MI, Bailey-Wilson J, Juo SH, Trent JM, Smith L, Brownstein MJ: Progressive juvenile-onset punctate cataracts caused by mutation of the gammaD-crystallin gene. Proc Natl Acad Sci USA 1999, 96:1008-1012.
67. Nandrot E, Slingsby C, Basak A, Cherif-Chefchaouni M, Benazzouz B, Hajaji Y, Boutayeb S, Gribouval O, Arbogast L, Berraho A, Abitbol M, Hilal L: Gamma-D crystallin gene (CRYGD) mutation causes autosomal dominant congenital cerulean cataracts. J Med Genet 2003, 40:262-267.
68. Plotnikova OV, Kondrashov FA, Vlasov PK, Grigorenko AP, Ginter EK, Rogaev EI: Conversion and compensatory evolution of the gammacrystallin genes and identification of a cataractogenic mutation that reverses the sequence of the human CRYGD gene to an ancestral state. Am J Hum Genet 2007, 81:32-43.
69. Kmoch S, Brynda J, Asfaw B, Bezouska K, Novak P, Rezacova P, Ondrova L, Filipec M, Sedlacek J, Elleder M: Link between a novel human gammaDcrystallin allele and a unique cataract phenotype explained by protein crystallography. Hum Mol Genet 2000, 9:1779-1786.
70. Wistow G, Sardarian L, Gan W, Wyatt MK: The human gene for gammaScrystallin: alternative transcripts and expressed sequences from the first intron. Mol Vis 2000, 6:79-84.
71. Sun H, Ma Z, Li Y, Liu B, Li Z, Ding X, Gao Y, Ma W, Tang X, Li X, Shen Y: Gamma-S crystallin gene (CRYGS) mutation causes dominant progressive cortical cataract in humans. J Med Genet 2005, 42:706-710.
72. Ray ME, Wistow G, Su YA, Meltzer PS, Trent JM: AIM1, a novel non-lens member of the betagamma-crystallin superfamily is associated with the control of tumorigenicity in human malignant melanoma. Proc Natl Acad Sci USA 1997, 94:3229-3234.
73. Aravind P, Wistow G, Sharma Y, Sankaranarayanan R: Exploring the limits of sequence and structure in a variant betagamma-crystallin domain of the protein absent in melanoma-1 (AIM1). J Mol Biol 2008, 381:509-518.
74. Liu SB, He YY, Zhang Y, Lee WH, Qian JQ, Lai R, Jin Y: A novel non-lens betagamma-crystallin and trefoil factor complex from amphibian skin and its functional implications. PLoS One 2008, 3:e1770.
75. Rabinowitz YS, Dong L, Wistow G: Gene expression profile studies of human keratoconus cornea for NEIBank: a novel cornea expressed gene and the absence of transcripts for aquaporin 5. Invest Ophthalmol Vis Sci 2005, 46:1239-1246.
76. Ray ME, Su YA, Meltzer PS, Trent JM: Isolation and characterization of genes associated with chromosome-6 mediated tumor suppression in human malignant melanoma. Oncogene 1996, 12:2527-2533.
77. Wistow G, Jaworski C, Rao PV: A non-lens member of the beta gammacrystallin superfamily in a vertebrate the amphibian Cynops. Exp Eye Res 1995, 61:637-639.
78. Takabatake T, Takahashi TC, Takeshima K: Cloning of an epidermis-specific Cynops cDNA from a neurula library. Develop Growth Differ 1992, 34:277-283.
79. Fan J, Dong L, Mishra S, Chen Y, Fitzgerald P, Wistow G: A role for gammaS-crystallin in the organization of actin and fiber cell maturation in the mouse lens. FEBS J 2012, 279:2892-2904.
80. Wyatt K, White HE, Wang L, Bateman OA, Slingsby C, Orlova EV, Wistow G: Lengsin is a survivor of an ancient family of class I glutamine synthetases re-engineered by evolution for a role in the vertebrate lens. Structure 2006, 14:1823-1834.
81. Wistow GJ, Mulders JW, de Jong WW: The enzyme lactate dehydrogenase as a structural protein in avian and crocodilian lenses. Nature 1987, 326:622-624.
82. Rao PV, Garrow TA, John F, Garland D, Millian NS, Zigler JS Jr: Betainehomocysteine methyltransferase is a developmentally regulated enzyme crystallin in rhesus monkey lens. J Biol Chem 1998, 273:30669-30674.
83. Wistow G, Kim H: Lens protein expression in mammals: taxon-specificity and the recruitment of crystallins. J Mol Evol 1991, 32:262-269.
84. Kim RY, Gasser R, Wistow GJ: Mu-crystallin is a mammalian homologue of Agrobacterium ornithine cyclodeaminase and is expressed in human retina. Proc Natl Acad Sci USA 1992, 89:9292-9296.
85. Segovia L, Horwitz J, Gasser R, Wistow G: Two roles for mu-crystallin: a lens structural protein in diurnal marsupials and a possible enzyme in mammalian retinas. Mol Vis 1997, 3:9.
86. Abe S, Katagiri T, Saito-Hisaminato A, Usami S, Inoue Y, Tsunoda T, Nakamura Y: Identification of CRYM as a candidate responsible for nonsyndromic deafness through cDNA microarray analysis of human cochlear and vestibular tissues. Am J Hum Genet 2003, 72:73-82.
87. Hallen A, Cooper AJ, Jamie JF, Haynes PA, Willows RD: Mammalian forebrain ketimine reductase identified as mu-crystallin; potential regulation by thyroid hormones. J Neurochem 2011, 118:379-387.
88. Vie MP, Evrard C, Osty J, Breton-Gilet A, Blanchet P, Pomerance M, Rouget P, Francon J, Blondeau JP: Purification, molecular cloning, and functional expression of the human nicodinamide-adenine dinucleotide phosphate-regulated thyroid hormone-binding protein. Mol Endocrinol 1997, 11:1728-1736.
89. Gonzalez P, Rao PV, Nunez SB, Zigler JS Jr: Evidence for independent recruitment of zeta-crystallin/quinone reductase (CRYZ) as a crystallin in camelids and hystricomorph rodents. Mol Biol Evol 1995, 12:773-781.
90. Rao PV, Krishna CM, Zigler JS: Identification and characterization of the enzymatic activity of zeta-crystallin from guinea pig lens. A novel NADPH:quinone oxidoreductase. J Biol Chem 1992, 267:96-102.
91. Mulders JW, Hendriks W, Blankesteijn WM, Bloemendal H, de Jong WW: Lambda-crystallin, a major rabbit lens protein, is related to hydroxyacylcoenzyme A dehydrogenases. J Biol Chem 1988, 263:15462-15466.
92. Wistow GJ, Lietman T, Williams LA, Stapel SO, de Jong WW, Horwitz J, Piatigorsky J: Tau-crystallin/alpha-enolase: one gene encodes both an enzyme and a lens structural protein. J Cell Biol 1988, 107:2729-2736.
93. Cvekl A, Yang Y, Chauhan BK, Cveklova K: Regulation of gene expression by Pax6 in ocular cells: a case of tissue-preferred expression of crystallins in lens. Int J Dev Biol 2004, 48:829-844.
94. Cvekl A, Duncan MK: Genetic and epigenetic mechanisms of gene regulation during lens development. Prog Retin Eye Res 2007, 26:555-597.
95. Sharon-Friling R, Richardson J, Sperbeck S, Lee D, Rauchman M, Maas R, Swaroop A, Wistow G: Lens-specific gene recruitment of zeta-crystallin through Pax6 Nrl-Maf, and brain suppressor sites. Mol Cell Biol 1998, 18:2067-2076.