Skip to main content
NCBI home page
The Journal of Cell Biology logoLink to The Journal of Cell Biology
. 1988 May 1;106(5):1635–1648. doi: 10.1083/jcb.106.5.1635

Expression of simple epithelial type cytokeratins in stratified epithelia as detected by immunolocalization and hybridization in situ

PMCID: PMC2115057  PMID: 2453518

Abstract

Multi-layered ("stratified") epithelia differ from one-layered ("simple") polar epithelia by various architectural and functional properties as well as by their cytoskeletal complements, notably a set of cytokeratins characteristic of stratified tissue. The simple epithelial cytokeratins 8 and 18 have so far not been detected in any stratified epithelium. Using specific monoclonal antibodies we have noted, in several but not all samples of stratified epithelia, including esophagus, tongue, exocervix, and vagina, positive immunocytochemical reactions for cytokeratins 8, 18, and 19 which in some regions were selective for the basal cell layer(s) but extended into suprabasal layers in others. In situ hybridization with different probes (riboprobes, synthetic oligonucleotides) for mRNAs of cytokeratin 8 on esophageal epithelium has shown, in extended regions, relatively strong reactivity for cytokeratin 8 mRNA in the basal cell layer. In contrast, probes to cytokeratin 18 have shown much weaker hybridization which, however, was rather evenly spread over basal and suprabasal strata. These results, which emphasize the importance of in situ hybridization in studies of gene expression in complex tissues, show that the genes encoding simple epithelial cytokeratins can be expressed in stratified epithelia. This suggests that continual expression of genes coding for simple epithelial cytokeratins is compatible with the formation of squamous stratified tissues and can occur, at least in basal cell layers, simultaneously with the synthesis of certain stratification-related cytokeratins. We also emphasize differences of expression and immunoreactivity of these cytokeratins between different samples and in different regions of the same stratified epithelium and discuss the results in relation to changes of cytokeratin expression during fetal development of stratified epithelia, in response to environmental factors and during the formation of squamous cell carcinomas.

Full Text

The Full Text of this article is available as a PDF (8.5 MB).

Selected References

These references are in PubMed. This may not be the complete list of references from this article.

  1. Achstätter T., Moll R., Anderson A., Kuhn C., Pitz S., Schwechheimer K., Franke W. W. Expression of glial filament protein (GFP) in nerve sheaths and non-neural cells re-examined using monoclonal antibodies, with special emphasis on the co-expression of GFP and cytokeratins in epithelial cells of human salivary gland and pleomorphic adenomas. Differentiation. 1986;31(3):206–227. doi: 10.1111/j.1432-0436.1986.tb00401.x. [DOI] [PubMed] [Google Scholar]
  2. Achtstätter T., Moll R., Moore B., Franke W. W. Cytokeratin polypeptide patterns of different epithelia of the human male urogenital tract: immunofluorescence and gel electrophoretic studies. J Histochem Cytochem. 1985 May;33(5):415–426. doi: 10.1177/33.5.2580881. [DOI] [PubMed] [Google Scholar]
  3. Banks-Schlegel S. P., Harris C. C. Aberrant expression of keratin proteins and cross-linked envelopes in human esophageal carcinomas. Cancer Res. 1984 Mar;44(3):1153–1157. [PubMed] [Google Scholar]
  4. Banks-Schlegel S. P., Harris C. C. Tissue-specific expression of keratin proteins in human esophageal and epidermal epithelium and their cultured keratinocytes. Exp Cell Res. 1983 Jul;146(2):271–280. doi: 10.1016/0014-4827(83)90129-5. [DOI] [PubMed] [Google Scholar]
  5. Banks-Schlegel S. P. Keratin alterations during embryonic epidermal differentiation: a presage of adult epidermal maturation. J Cell Biol. 1982 Jun;93(3):551–559. doi: 10.1083/jcb.93.3.551. [DOI] [PMC free article] [PubMed] [Google Scholar]
  6. Banks-Schlegel S. P., Quintero J. Growth and differentiation of human esophageal carcinoma cell lines. Cancer Res. 1986 Jan;46(1):250–258. [PubMed] [Google Scholar]
  7. Bártek J., Bártková J., Taylor-Papadimitriou J., Rejthar A., Kovarík J., Lukás Z., Vojtesek B. Differential expression of keratin 19 in normal human epithelial tissues revealed by monospecific monoclonal antibodies. Histochem J. 1986 Oct;18(10):565–575. doi: 10.1007/BF01675198. [DOI] [PubMed] [Google Scholar]
  8. Cooper D., Schermer A., Sun T. T. Classification of human epithelia and their neoplasms using monoclonal antibodies to keratins: strategies, applications, and limitations. Lab Invest. 1985 Mar;52(3):243–256. [PubMed] [Google Scholar]
  9. Cooper D., Sun T. T. Monoclonal antibody analysis of bovine epithelial keratins. Specific pairs as defined by coexpression. J Biol Chem. 1986 Apr 5;261(10):4646–4654. [PubMed] [Google Scholar]
  10. Danto S. I., Fischman D. A. Immunocytochemical analysis of intermediate filaments in embryonic heart cells with monoclonal antibodies to desmin. J Cell Biol. 1984 Jun;98(6):2179–2191. doi: 10.1083/jcb.98.6.2179. [DOI] [PMC free article] [PubMed] [Google Scholar]
  11. Debus E., Moll R., Franke W. W., Weber K., Osborn M. Immunohistochemical distinction of human carcinomas by cytokeratin typing with monoclonal antibodies. Am J Pathol. 1984 Jan;114(1):121–130. [PMC free article] [PubMed] [Google Scholar]
  12. Debus E., Weber K., Osborn M. Monoclonal cytokeratin antibodies that distinguish simple from stratified squamous epithelia: characterization on human tissues. EMBO J. 1982;1(12):1641–1647. doi: 10.1002/j.1460-2075.1982.tb01367.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
  13. Dixon I. S., Stanley M. A. Immunofluorescent studies of human cervical epithelia in vivo and in vitro using antibodies against specific keratin components. Mol Biol Med. 1984 Feb;2(1):37–51. [PubMed] [Google Scholar]
  14. Dulbecco R., Allen R., Okada S., Bowman M. Functional changes of intermediate filaments in fibroblastic cells revealed by a monoclonal antibody. Proc Natl Acad Sci U S A. 1983 Apr;80(7):1915–1918. doi: 10.1073/pnas.80.7.1915. [DOI] [PMC free article] [PubMed] [Google Scholar]
  15. Eichner R., Sun T. T., Aebi U. The role of keratin subfamilies and keratin pairs in the formation of human epidermal intermediate filaments. J Cell Biol. 1986 May;102(5):1767–1777. doi: 10.1083/jcb.102.5.1767. [DOI] [PMC free article] [PubMed] [Google Scholar]
  16. Franke W. W., Appelhans B., Schmid E., Freudenstein C., Osborn M., Weber K. Identification and characterization of epithelial cells in mammalian tissues by immunofluorescence microscopy using antibodies to prekeratin. Differentiation. 1979;15(1):7–25. doi: 10.1111/j.1432-0436.1979.tb01030.x. [DOI] [PubMed] [Google Scholar]
  17. Franke W. W., Grund C., Kuhn C., Lehto V. P., Virtanen I. Transient change of organization of vimentin filaments during mitosis as demonstrated by a monoclonal antibody. Exp Cell Res. 1984 Oct;154(2):567–580. doi: 10.1016/0014-4827(84)90181-2. [DOI] [PubMed] [Google Scholar]
  18. Franke W. W., Schiller D. L., Moll R., Winter S., Schmid E., Engelbrecht I., Denk H., Krepler R., Platzer B. Diversity of cytokeratins. Differentiation specific expression of cytokeratin polypeptides in epithelial cells and tissues. J Mol Biol. 1981 Dec 25;153(4):933–959. doi: 10.1016/0022-2836(81)90460-5. [DOI] [PubMed] [Google Scholar]
  19. Franke W. W., Schmid E., Osborn M., Weber K. Different intermediate-sized filaments distinguished by immunofluorescence microscopy. Proc Natl Acad Sci U S A. 1978 Oct;75(10):5034–5038. doi: 10.1073/pnas.75.10.5034. [DOI] [PMC free article] [PubMed] [Google Scholar]
  20. Franke W. W., Schmid E., Schiller D. L., Winter S., Jarasch E. D., Moll R., Denk H., Jackson B. W., Illmensee K. Differentiation-related patterns of expression of proteins of intermediate-size filaments in tissues and cultured cells. Cold Spring Harb Symp Quant Biol. 1982;46(Pt 1):431–453. doi: 10.1101/sqb.1982.046.01.041. [DOI] [PubMed] [Google Scholar]
  21. Franke W. W., Schmid E., Wellsteed J., Grund C., Gigi O., Geiger B. Change of cytokeratin filament organization during the cell cycle: selective masking of an immunologic determinant in interphase PtK2 cells. J Cell Biol. 1983 Oct;97(4):1255–1260. doi: 10.1083/jcb.97.4.1255. [DOI] [PMC free article] [PubMed] [Google Scholar]
  22. Franke W. W., Weber K., Osborn M., Schmid E., Freudenstein C. Antibody to prekeratin. Decoration of tonofilament like arrays in various cells of epithelial character. Exp Cell Res. 1978 Oct 15;116(2):429–445. doi: 10.1016/0014-4827(78)90466-4. [DOI] [PubMed] [Google Scholar]
  23. Franke W. W., Winter S., Schmid E., Söllner P., Hämmerling G., Achtstätter T. Monoclonal cytokeratin antibody recognizing a heterotypic complex: immunological probing of conformational states of cytoskeletal proteins in filaments and in solution. Exp Cell Res. 1987 Nov;173(1):17–37. doi: 10.1016/0014-4827(87)90328-4. [DOI] [PubMed] [Google Scholar]
  24. Franke W. W., Winter S., von Overbeck J., Gudat F., Heitz P. U., Stähli C. Identification of the conserved, conformation-dependent cytokeratin epitope recognized by monoclonal antibody (lu-5). Virchows Arch A Pathol Anat Histopathol. 1987;411(2):137–147. doi: 10.1007/BF00712737. [DOI] [PubMed] [Google Scholar]
  25. Fuchs E. V., Coppock S. M., Green H., Cleveland D. W. Two distinct classes of keratin genes and their evolutionary significance. Cell. 1981 Nov;27(1 Pt 2):75–84. doi: 10.1016/0092-8674(81)90362-7. [DOI] [PubMed] [Google Scholar]
  26. Fuchs E., Green H. Changes in keratin gene expression during terminal differentiation of the keratinocyte. Cell. 1980 Apr;19(4):1033–1042. doi: 10.1016/0092-8674(80)90094-x. [DOI] [PubMed] [Google Scholar]
  27. Fuchs E., Green H. Multiple keratins of cultured human epidermal cells are translated from different mRNA molecules. Cell. 1979 Jul;17(3):573–582. doi: 10.1016/0092-8674(79)90265-4. [DOI] [PubMed] [Google Scholar]
  28. Fuchs E., Green H. Regulation of terminal differentiation of cultured human keratinocytes by vitamin A. Cell. 1981 Sep;25(3):617–625. doi: 10.1016/0092-8674(81)90169-0. [DOI] [PubMed] [Google Scholar]
  29. Gilfix B. M., Eckert R. L. Coordinate control by vitamin A of keratin gene expression in human keratinocytes. J Biol Chem. 1985 Nov 15;260(26):14026–14029. [PubMed] [Google Scholar]
  30. Gown A. M., Vogel A. M. Monoclonal antibodies to human intermediate filament proteins. II. Distribution of filament proteins in normal human tissues. Am J Pathol. 1984 Feb;114(2):309–321. [PMC free article] [PubMed] [Google Scholar]
  31. Grace M. P., Kim K. H., True L. D., Fuchs E. Keratin expression in normal esophageal epithelium and squamous cell carcinoma of the esophagus. Cancer Res. 1985 Feb;45(2):841–846. [PubMed] [Google Scholar]
  32. Hanukoglu I., Fuchs E. The cDNA sequence of a Type II cytoskeletal keratin reveals constant and variable structural domains among keratins. Cell. 1983 Jul;33(3):915–924. doi: 10.1016/0092-8674(83)90034-x. [DOI] [PubMed] [Google Scholar]
  33. Hatzfeld M., Franke W. W. Pair formation and promiscuity of cytokeratins: formation in vitro of heterotypic complexes and intermediate-sized filaments by homologous and heterologous recombinations of purified polypeptides. J Cell Biol. 1985 Nov;101(5 Pt 1):1826–1841. doi: 10.1083/jcb.101.5.1826. [DOI] [PMC free article] [PubMed] [Google Scholar]
  34. Hazan R., Denk H., Franke W. W., Lackinger E., Schiller D. L. Change of cytokeratin organization during development of Mallory bodies as revealed by a monoclonal antibody. Lab Invest. 1986 May;54(5):543–553. [PubMed] [Google Scholar]
  35. Heid H. W., Werner E., Franke W. W. The complement of native alpha-keratin polypeptides of hair-forming cells: a subset of eight polypeptides that differ from epithelial cytokeratins. Differentiation. 1986;32(2):101–119. doi: 10.1111/j.1432-0436.1986.tb00562.x. [DOI] [PubMed] [Google Scholar]
  36. Hopwood D., Coghill G., Sanders D. S. Human oesophageal submucosal glands. Their detection mucin, enzyme and secretory protein content. Histochemistry. 1986;86(1):107–112. doi: 10.1007/BF00492353. [DOI] [PubMed] [Google Scholar]
  37. Jorcano J. L., Rieger M., Franz J. K., Schiller D. L., Moll R., Franke W. W. Identification of two types of keratin polypeptides within the acidic cytokeratin subfamily I. J Mol Biol. 1984 Oct 25;179(2):257–281. doi: 10.1016/0022-2836(84)90468-6. [DOI] [PubMed] [Google Scholar]
  38. Karsten U., Papsdorf G., Roloff G., Stolley P., Abel H., Walther I., Weiss H. Monoclonal anti-cytokeratin antibody from a hybridoma clone generated by electrofusion. Eur J Cancer Clin Oncol. 1985 Jun;21(6):733–740. doi: 10.1016/0277-5379(85)90271-8. [DOI] [PubMed] [Google Scholar]
  39. Kemler R., Brûlet P., Schnebelen M. T., Gaillard J., Jacob F. Reactivity of monoclonal antibodies against intermediate filament proteins during embryonic development. J Embryol Exp Morphol. 1981 Aug;64:45–60. [PubMed] [Google Scholar]
  40. Kopan R., Traska G., Fuchs E. Retinoids as important regulators of terminal differentiation: examining keratin expression in individual epidermal cells at various stages of keratinization. J Cell Biol. 1987 Jul;105(1):427–440. doi: 10.1083/jcb.105.1.427. [DOI] [PMC free article] [PubMed] [Google Scholar]
  41. Lane E. B., Bártek J., Purkis P. E., Leigh I. M. Keratin antigens in differentiating skin. Ann N Y Acad Sci. 1985;455:241–258. doi: 10.1111/j.1749-6632.1985.tb50415.x. [DOI] [PubMed] [Google Scholar]
  42. Lane E. B. Monoclonal antibodies provide specific intramolecular markers for the study of epithelial tonofilament organization. J Cell Biol. 1982 Mar;92(3):665–673. doi: 10.1083/jcb.92.3.665. [DOI] [PMC free article] [PubMed] [Google Scholar]
  43. Lavker R. M., Sun T. T. Epidermal stem cells. J Invest Dermatol. 1983 Jul;81(1 Suppl):121s–127s. doi: 10.1111/1523-1747.ep12540880. [DOI] [PubMed] [Google Scholar]
  44. Lazarides E., Granger B. L., Gard D. L., O'Connor C. M., Breckler J., Price M., Danto S. I. Desmin- and vimentin-containing filaments and their role inthe assembly of the Z disk in muscle cells. Cold Spring Harb Symp Quant Biol. 1982;46(Pt 1):351–378. doi: 10.1101/sqb.1982.046.01.036. [DOI] [PubMed] [Google Scholar]
  45. Lee L. D., Baden H. P. Organisation of the polypeptide chains in mammalian keratin. Nature. 1976 Nov 25;264(5584):377–379. doi: 10.1038/264377a0. [DOI] [PubMed] [Google Scholar]
  46. Leube R. E., Bader B. L., Bosch F. X., Zimbelmann R., Achtstaetter T., Franke W. W. Molecular characterization and expression of the stratification-related cytokeratins 4 and 15. J Cell Biol. 1988 Apr;106(4):1249–1261. doi: 10.1083/jcb.106.4.1249. [DOI] [PMC free article] [PubMed] [Google Scholar]
  47. Leube R. E., Bosch F. X., Romano V., Zimbelmann R., Höfler H., Franke W. W. Cytokeratin expression in simple epithelia. III. Detection of mRNAs encoding human cytokeratins nos. 8 and 18 in normal and tumor cells by hybridization with cDNA sequences in vitro and in situ. Differentiation. 1986;33(1):69–85. doi: 10.1111/j.1432-0436.1986.tb00412.x. [DOI] [PubMed] [Google Scholar]
  48. Lynch M. H., O'Guin W. M., Hardy C., Mak L., Sun T. T. Acidic and basic hair/nail ("hard") keratins: their colocalization in upper cortical and cuticle cells of the human hair follicle and their relationship to "soft" keratins. J Cell Biol. 1986 Dec;103(6 Pt 2):2593–2606. doi: 10.1083/jcb.103.6.2593. [DOI] [PMC free article] [PubMed] [Google Scholar]
  49. Magin T. M., Jorcano J. L., Franke W. W. Cytokeratin expression in simple epithelia. II. cDNA cloning and sequence characteristics of bovine cytokeratin A (no. 8). Differentiation. 1986;30(3):254–264. doi: 10.1111/j.1432-0436.1986.tb00788.x. [DOI] [PubMed] [Google Scholar]
  50. Makin C. A., Bobrow L. G., Bodmer W. F. Monoclonal antibody to cytokeratin for use in routine histopathology. J Clin Pathol. 1984 Sep;37(9):975–983. doi: 10.1136/jcp.37.9.975. [DOI] [PMC free article] [PubMed] [Google Scholar]
  51. Marchuk D., McCrohon S., Fuchs E. Complete sequence of a gene encoding a human type I keratin: sequences homologous to enhancer elements in the regulatory region of the gene. Proc Natl Acad Sci U S A. 1985 Mar;82(6):1609–1613. doi: 10.1073/pnas.82.6.1609. [DOI] [PMC free article] [PubMed] [Google Scholar]
  52. Milstone L. M. Isolation and characterization of two polypeptides that form intermediate filaments in bovine esophageal epithelium. J Cell Biol. 1981 Feb;88(2):317–322. doi: 10.1083/jcb.88.2.317. [DOI] [PMC free article] [PubMed] [Google Scholar]
  53. Moll R., Franke W. W., Schiller D. L., Geiger B., Krepler R. The catalog of human cytokeratins: patterns of expression in normal epithelia, tumors and cultured cells. Cell. 1982 Nov;31(1):11–24. doi: 10.1016/0092-8674(82)90400-7. [DOI] [PubMed] [Google Scholar]
  54. Moll R., Krepler R., Franke W. W. Complex cytokeratin polypeptide patterns observed in certain human carcinomas. Differentiation. 1983;23(3):256–269. doi: 10.1111/j.1432-0436.1982.tb01291.x. [DOI] [PubMed] [Google Scholar]
  55. Moll R., Levy R., Czernobilsky B., Hohlweg-Majert P., Dallenbach-Hellweg G., Franke W. W. Cytokeratins of normal epithelia and some neoplasms of the female genital tract. Lab Invest. 1983 Nov;49(5):599–610. [PubMed] [Google Scholar]
  56. Moll R., Moll I., Franke W. W. Identification of Merkel cells in human skin by specific cytokeratin antibodies: changes of cell density and distribution in fetal and adult plantar epidermis. Differentiation. 1984;28(2):136–154. doi: 10.1111/j.1432-0436.1984.tb00277.x. [DOI] [PubMed] [Google Scholar]
  57. Moll R., Moll I., Wiest W. Changes in the pattern of cytokeratin polypeptides in epidermis and hair follicles during skin development in human fetuses. Differentiation. 1982;23(2):170–178. doi: 10.1111/j.1432-0436.1982.tb01280.x. [DOI] [PubMed] [Google Scholar]
  58. Morris A., Steinberg M. L., Defendi V. Keratin gene expression in simian virus 40-transformed human keratinocytes. Proc Natl Acad Sci U S A. 1985 Dec;82(24):8498–8502. doi: 10.1073/pnas.82.24.8498. [DOI] [PMC free article] [PubMed] [Google Scholar]
  59. Nagle R. B., Moll R., Weidauer H., Nemetschek H., Franke W. W. Different patterns of cytokeratin expression in the normal epithelia of the upper respiratory tract. Differentiation. 1985;30(2):130–140. doi: 10.1111/j.1432-0436.1985.tb00524.x. [DOI] [PubMed] [Google Scholar]
  60. Ouhayoun J. P., Gosselin F., Forest N., Winter S., Franke W. W. Cytokeratin patterns of human oral epithelia: differences in cytokeratin synthesis in gingival epithelium and the adjacent alveolar mucosa. Differentiation. 1985;30(2):123–129. doi: 10.1111/j.1432-0436.1985.tb00523.x. [DOI] [PubMed] [Google Scholar]
  61. Pallesen G., Nielsen S., Celis J. E. Characterization of a monoclonal antibody (BG3C8) that reacts with basal cells of stratified epithelia. Histopathology. 1987 Jun;11(6):591–601. doi: 10.1111/j.1365-2559.1987.tb02669.x. [DOI] [PubMed] [Google Scholar]
  62. Quinlan R. A., Cohlberg J. A., Schiller D. L., Hatzfeld M., Franke W. W. Heterotypic tetramer (A2D2) complexes of non-epidermal keratins isolated from cytoskeletons of rat hepatocytes and hepatoma cells. J Mol Biol. 1984 Sep 15;178(2):365–388. doi: 10.1016/0022-2836(84)90149-9. [DOI] [PubMed] [Google Scholar]
  63. Ramaekers F., Huysmans A., Moesker O., Kant A., Jap P., Herman C., Vooijs P. Monoclonal antibody to keratin filaments, specific for glandular epithelia and their tumors. Use in surgical pathology. Lab Invest. 1983 Sep;49(3):353–361. [PubMed] [Google Scholar]
  64. Ramaekers F., Huysmans A., Schaart G., Moesker O., Vooijs P. Tissue distribution of keratin 7 as monitored by a monoclonal antibody. Exp Cell Res. 1987 May;170(1):235–249. doi: 10.1016/0014-4827(87)90133-9. [DOI] [PubMed] [Google Scholar]
  65. Regauer S., Franke W. W., Virtanen I. Intermediate filament cytoskeleton of amnion epithelium and cultured amnion epithelial cells: expression of epidermal cytokeratins in cells of a simple epithelium. J Cell Biol. 1985 Apr;100(4):997–1009. doi: 10.1083/jcb.100.4.997. [DOI] [PMC free article] [PubMed] [Google Scholar]
  66. Rentrop M., Knapp B., Winter H., Schweizer J. Differential localization of distinct keratin mRNA-species in mouse tongue epithelium by in situ hybridization with specific cDNA probes. J Cell Biol. 1986 Dec;103(6 Pt 2):2583–2591. doi: 10.1083/jcb.103.6.2583. [DOI] [PMC free article] [PubMed] [Google Scholar]
  67. Romano V., Hatzfeld M., Magin T. M., Zimbelmann R., Franke W. W., Maier G., Ponstingl H. Cytokeratin expression in simple epithelia. I. Identification of mRNA coding for human cytokeratin no. 18 by a cDNA clone. Differentiation. 1986;30(3):244–253. doi: 10.1111/j.1432-0436.1986.tb00787.x. [DOI] [PubMed] [Google Scholar]
  68. Rupniak H. T., Rowlatt C., Lane E. B., Steele J. G., Trejdosiewicz L. K., Laskiewicz B., Povey S., Hill B. T. Characteristics of four new human cell lines derived from squamous cell carcinomas of the head and neck. J Natl Cancer Inst. 1985 Oct;75(4):621–635. [PubMed] [Google Scholar]
  69. Saurat J. H., Didierjean L. The epidermal Merkel cell is an epithelial cell. Dermatologica. 1984;169(3):117–120. doi: 10.1159/000249584. [DOI] [PubMed] [Google Scholar]
  70. Schiller D. L., Franke W. W., Geiger B. A subfamily of relatively large and basic cytokeratin polypeptides as defined by peptide mapping is represented by one or several polypeptides in epithelial cells. EMBO J. 1982;1(6):761–769. doi: 10.1002/j.1460-2075.1982.tb01243.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
  71. Schweizer J., Kinjo M., Fürstenberger G., Winter H. Sequential expression of mRNA-encoded keratin sets in neonatal mouse epidermis: basal cells with properties of terminally differentiating cells. Cell. 1984 May;37(1):159–170. doi: 10.1016/0092-8674(84)90311-8. [DOI] [PubMed] [Google Scholar]
  72. Steinert P. M., Idler W. W., Zimmerman S. B. Self-assembly of bovine epidermal keratin filaments in vitro. J Mol Biol. 1976 Dec 15;108(3):547–567. doi: 10.1016/s0022-2836(76)80136-2. [DOI] [PubMed] [Google Scholar]
  73. Steinert P. M., Steven A. C., Roop D. R. The molecular biology of intermediate filaments. Cell. 1985 Sep;42(2):411–420. doi: 10.1016/0092-8674(85)90098-4. [DOI] [PubMed] [Google Scholar]
  74. Sun T. T., Green H. Immunofluorescent staining of keratin fibers in cultured cells. Cell. 1978 Jul;14(3):469–476. doi: 10.1016/0092-8674(78)90233-7. [DOI] [PubMed] [Google Scholar]
  75. Sun T. T., Shih C., Green H. Keratin cytoskeletons in epithelial cells of internal organs. Proc Natl Acad Sci U S A. 1979 Jun;76(6):2813–2817. doi: 10.1073/pnas.76.6.2813. [DOI] [PMC free article] [PubMed] [Google Scholar]
  76. Sun T. T., Tseng S. C., Huang A. J., Cooper D., Schermer A., Lynch M. H., Weiss R., Eichner R. Monoclonal antibody studies of mammalian epithelial keratins: a review. Ann N Y Acad Sci. 1985;455:307–329. doi: 10.1111/j.1749-6632.1985.tb50419.x. [DOI] [PubMed] [Google Scholar]
  77. Taylor-Papadimitriou J., Purkis P., Lane E. B., McKay I. A., Chang S. E. Effects of SV40 transformation on the cytoskeleton and behavioural properties of human keratinocytes. Cell Differ. 1982 May;11(3):169–180. doi: 10.1016/0045-6039(82)90008-2. [DOI] [PubMed] [Google Scholar]
  78. Tseng S. C., Jarvinen M. J., Nelson W. G., Huang J. W., Woodcock-Mitchell J., Sun T. T. Correlation of specific keratins with different types of epithelial differentiation: monoclonal antibody studies. Cell. 1982 Sep;30(2):361–372. doi: 10.1016/0092-8674(82)90234-3. [DOI] [PubMed] [Google Scholar]
  79. Tyner A. L., Fuchs E. Evidence for posttranscriptional regulation of the keratins expressed during hyperproliferation and malignant transformation in human epidermis. J Cell Biol. 1986 Nov;103(5):1945–1955. doi: 10.1083/jcb.103.5.1945. [DOI] [PMC free article] [PubMed] [Google Scholar]
  80. Van Muijen G. N., Warnaar S. O., Ponec M. Differentiation-related changes of cytokeratin expression in cultured keratinocytes and in fetal, newborn, and adult epidermis. Exp Cell Res. 1987 Aug;171(2):331–345. doi: 10.1016/0014-4827(87)90166-2. [DOI] [PubMed] [Google Scholar]
  81. Viac J., Reano A., Brochier J., Staquet M. J., Thivolet J. Reactivity pattern of a monoclonal antikeratin antibody (KL1). J Invest Dermatol. 1983 Oct;81(4):351–354. doi: 10.1111/1523-1747.ep12519941. [DOI] [PubMed] [Google Scholar]
  82. Virtanen I., Miettinen M., Lehto V. P., Kariniemi A. L., Paasivuo R. Diagnostic application of monoclonal antibodies to intermediate filaments. Ann N Y Acad Sci. 1985;455:635–648. doi: 10.1111/j.1749-6632.1985.tb50441.x. [DOI] [PubMed] [Google Scholar]
  83. Wild G. A., Mischke D. Variation and frequency of cytokeratin polypeptide patterns in human squamous non-keratinizing epithelium. Exp Cell Res. 1986 Jan;162(1):114–126. doi: 10.1016/0014-4827(86)90430-1. [DOI] [PubMed] [Google Scholar]
  84. Woodcock-Mitchell J., Eichner R., Nelson W. G., Sun T. T. Immunolocalization of keratin polypeptides in human epidermis using monoclonal antibodies. J Cell Biol. 1982 Nov;95(2 Pt 1):580–588. doi: 10.1083/jcb.95.2.580. [DOI] [PMC free article] [PubMed] [Google Scholar]
  85. Woods E. F. The number of polypeptide chains in the rod domain of bovine epidermal keratin. Biochem Int. 1983 Dec;7(6):769–774. [PubMed] [Google Scholar]
  86. Wu R., Wu M. M. Effects of retinoids on human bronchial epithelial cells: differential regulation of hyaluronate synthesis and keratin protein synthesis. J Cell Physiol. 1986 Apr;127(1):73–82. doi: 10.1002/jcp.1041270110. [DOI] [PubMed] [Google Scholar]
  87. Wu Y. J., Parker L. M., Binder N. E., Beckett M. A., Sinard J. H., Griffiths C. T., Rheinwald J. G. The mesothelial keratins: a new family of cytoskeletal proteins identified in cultured mesothelial cells and nonkeratinizing epithelia. Cell. 1982 Dec;31(3 Pt 2):693–703. doi: 10.1016/0092-8674(82)90324-5. [DOI] [PubMed] [Google Scholar]
  88. Wu Y. J., Rheinwald J. G. A new small (40 kd) keratin filament protein made by some cultured human squamous cell carcinomas. Cell. 1981 Sep;25(3):627–635. doi: 10.1016/0092-8674(81)90170-7. [DOI] [PubMed] [Google Scholar]
  89. van Muijen G. N., Ruiter D. J., Franke W. W., Achtstätter T., Haasnoot W. H., Ponec M., Warnaar S. O. Cell type heterogeneity of cytokeratin expression in complex epithelia and carcinomas as demonstrated by monoclonal antibodies specific for cytokeratins nos. 4 and 13. Exp Cell Res. 1986 Jan;162(1):97–113. doi: 10.1016/0014-4827(86)90429-5. [DOI] [PubMed] [Google Scholar]
  90. van Muijen G. N., Ruiter D. J., Ponec M., Huiskens-van der, Mey C., Warnaar S. O. Monoclonal antibodies with different specificities against cytokeratins. An immunohistochemical study of normal tissues and tumors. Am J Pathol. 1984 Jan;114(1):9–17. [PMC free article] [PubMed] [Google Scholar]

Articles from The Journal of Cell Biology are provided here courtesy of The Rockefeller University Press

RESOURCES