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Gene Review

Wap  -  whey acidic protein

Mus musculus

Synonyms: WAP, Whey acidic protein
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Disease relevance of Wap


High impact information on Wap


Chemical compound and disease context of Wap


Biological context of Wap


Anatomical context of Wap


Associations of Wap with chemical compounds


Regulatory relationships of Wap


Other interactions of Wap

  • The increase of Stat5 expression during pregnancy coincides with the activation of the WAP gene [24].
  • Non-virgin WAP-TGFalpha transgenic mice displayed accelerated mammary development during pregnancy, delayed post-parturient mammary involution, a progressive increase in the number of hyperplastic alveolar nodules (HANs), and development of mammary carcinoma with a mean latency of 9 months [1].
  • Even though the CEL and WAP genes are induced at the same time during mammary gland differentiation, their regulation is different with respect to dependence on lactogenic hormones [25].
  • We generated transgenic mice that express autoactivated isoforms of the matrix metalloproteinase stromelysin-1, under the control of the whey acidic protein gene promoter, to examine the effect of inappropriate expression of this enzyme [26].
  • Some but not all of these NFI isoforms synergistically activate WAP gene transcription in cooperation with GR and STAT5, as determined using transient cotransfection assays in JEG-3 cells [27].

Analytical, diagnostic and therapeutic context of Wap

  • In the present study, we investigated, by means of the reverse transcription polymerase chain reaction (RT-PCR), the patterns of expression of endogenous WAP genes in tissues of normal mice and in transgenic mice carrying hGH gene coupled to the mWAP promoter sequence [14].
  • The regulation of the Wap gene differs from that of other milk protein genes, with one consequence being that little or no Wap expression is detectable in cell culture [18].
  • The WAP/bTAP transcript was detected in RNA isolated from mammary tissue of transgenic females. bTAP was purified to homogeneity from milk via acid precipitation, reverse-phase HPLC, and ion-exchange chromatography [28].
  • By using a whole mount organ culture system to study the differentiation potential of the mammary epithelium, we observed a reduced number of fully developed alveoli and a decrease in whey acidic protein expression [29].
  • Transplantation of WAP-TGF beta 1 mammary gland into nonpregnant hosts revealed that transgenic implants, even those from young postpubertal virgin females, had a diminished ability to repopulate epithelium-free mammary fat pads [30].


  1. Transforming growth factor alpha- and c-myc-induced mammary carcinogenesis in transgenic mice. Rose-Hellekant, T.A., Sandgren, E.P. Oncogene (2000) [Pubmed]
  2. Mammary tumor induction in transgenic mice expressing an RNA-binding protein. Tessier, C.R., Doyle, G.A., Clark, B.A., Pitot, H.C., Ross, J. Cancer Res. (2004) [Pubmed]
  3. Targeted expression of HGF/SF in mouse mammary epithelium leads to metastatic adenosquamous carcinomas through the activation of multiple signal transduction pathways. Gallego, M.I., Bierie, B., Hennighausen, L. Oncogene (2003) [Pubmed]
  4. Whey acidic protein extrinsically expressed from the mouse mammary tumor virus long terminal repeat results in hyperplasia of the coagulation gland epithelium and impaired mammary development. Hennighausen, L., McKnight, R., Burdon, T., Baik, M., Wall, R.J., Smith, G.H. Cell Growth Differ. (1994) [Pubmed]
  5. Ha-ras and c-myc oncogene expression interferes with morphological and functional differentiation of mammary epithelial cells in single and double transgenic mice. Andres, A.C., van der Valk, M.A., Schönenberger, C.A., Flückiger, F., LeMeur, M., Gerlinger, P., Groner, B. Genes Dev. (1988) [Pubmed]
  6. Essential functions of p21-activated kinase 1 in morphogenesis and differentiation of mammary glands. Wang, R.A., Vadlamudi, R.K., Bagheri-Yarmand, R., Beuvink, I., Hynes, N.E., Kumar, R. J. Cell Biol. (2003) [Pubmed]
  7. Perturbation of beta1-integrin function alters the development of murine mammary gland. Faraldo, M.M., Deugnier, M.A., Lukashev, M., Thiery, J.P., Glukhova, M.A. EMBO J. (1998) [Pubmed]
  8. Extracellular matrix regulates whey acidic protein gene expression by suppression of TGF-alpha in mouse mammary epithelial cells: studies in culture and in transgenic mice. Lin, C.Q., Dempsey, P.J., Coffey, R.J., Bissell, M.J. J. Cell Biol. (1995) [Pubmed]
  9. Chemoprevention of mammary carcinogenesis in a transgenic mouse model by alpha-difluoromethylornithine (DFMO) in the diet is associated with decreased cyclin D1 activity. Li, M., Ren, S., Tilli, M.T., Flaws, J.A., Lubet, R., Grubbs, C.J., Furth, P.A. Oncogene (2003) [Pubmed]
  10. Nuclear factor I and mammary gland factor (STAT5) play a critical role in regulating rat whey acidic protein gene expression in transgenic mice. Li, S., Rosen, J.M. Mol. Cell. Biol. (1995) [Pubmed]
  11. Targeted c-myc gene expression in mammary glands of transgenic mice induces mammary tumours with constitutive milk protein gene transcription. Schoenenberger, C.A., Andres, A.C., Groner, B., van der Valk, M., LeMeur, M., Gerlinger, P. EMBO J. (1988) [Pubmed]
  12. Modulation of secreted proteins of mouse mammary epithelial cells by the collagenous substrata. Lee, E.Y., Parry, G., Bissell, M.J. J. Cell Biol. (1984) [Pubmed]
  13. Cloning, transcription and chromosomal localization of the porcine whey acidic protein gene and its expression in HC11 cell line. Rival, S., Attal, J., Delville-Giraud, C., Yerle, M., Laffont, P., Rogel-Gaillard, C., Houdebine, L. Gene (2001) [Pubmed]
  14. Expression of whey acidic protein (WAP) genes in tissues other than the mammary gland in normal and transgenic mice expressing mWAP/hGH fusion gene. Wen, J., Kawamata, Y., Tojo, H., Tanaka, S., Tachi, C. Mol. Reprod. Dev. (1995) [Pubmed]
  15. Inhibition of myc-dependent breast tumor formation in transgenic mice. Cui, W., Gusterson, B.A., Clark, A.J. Breast Cancer Res. Treat. (2002) [Pubmed]
  16. Ha-ras oncogene expression directed by a milk protein gene promoter: tissue specificity, hormonal regulation, and tumor induction in transgenic mice. Andres, A.C., Schönenberger, C.A., Groner, B., Hennighausen, L., LeMeur, M., Gerlinger, P. Proc. Natl. Acad. Sci. U.S.A. (1987) [Pubmed]
  17. SV40 T-antigen induces breast cancer formation with a high efficiency in lactating and virgin WAP-SV-T transgenic animals but with a low efficiency in ovariectomized animals. Santarelli, R., Tzeng, Y.J., Zimmermann, C., Guhl, E., Graessmann, A. Oncogene (1996) [Pubmed]
  18. Induction of the endogenous whey acidic protein (Wap) gene and a Wap-myc hybrid gene in primary murine mammary organoids. Schoenenberger, C.A., Zuk, A., Groner, B., Jones, W., Andres, A.C. Dev. Biol. (1990) [Pubmed]
  19. Promotion of angiogenesis by ps20 in the differential reactive stroma prostate cancer xenograft model. McAlhany, S.J., Ressler, S.J., Larsen, M., Tuxhorn, J.A., Yang, F., Dang, T.D., Rowley, D.R. Cancer Res. (2003) [Pubmed]
  20. Expression of a whey acidic protein transgene during mammary development. Evidence for different mechanisms of regulation during pregnancy and lactation. Burdon, T., Sankaran, L., Wall, R.J., Spencer, M., Hennighausen, L. J. Biol. Chem. (1991) [Pubmed]
  21. Activating transcription factor 4 overexpression inhibits proliferation and differentiation of mammary epithelium resulting in impaired lactation and accelerated involution. Bagheri-Yarmand, R., Vadlamudi, R.K., Kumar, R. J. Biol. Chem. (2003) [Pubmed]
  22. Conditional deletion of the bcl-x gene from mouse mammary epithelium results in accelerated apoptosis during involution but does not compromise cell function during lactation. Walton, K.D., Wagner, K.U., Rucker, E.B., Shillingford, J.M., Miyoshi, K., Hennighausen, L. Mech. Dev. (2001) [Pubmed]
  23. Kit and PDGFR-alpha activities are necessary for Notch4/Int3-induced tumorigenesis. Raafat, A., Zoltan-Jones, A., Strizzi, L., Bargo, S., Kimura, K., Salomon, D., Callahan, R. Oncogene (2007) [Pubmed]
  24. Cloning and expression of Stat5 and an additional homologue (Stat5b) involved in prolactin signal transduction in mouse mammary tissue. Liu, X., Robinson, G.W., Gouilleux, F., Groner, B., Hennighausen, L. Proc. Natl. Acad. Sci. U.S.A. (1995) [Pubmed]
  25. Studies of the regulation of the mouse carboxyl ester lipase gene in mammary gland. Kannius-Janson, M., Lidberg, U., Hultén, K., Gritli-Linde, A., Bjursell, G., Nilsson, J. Biochem. J. (1998) [Pubmed]
  26. Targeted expression of stromelysin-1 in mammary gland provides evidence for a role of proteinases in branching morphogenesis and the requirement for an intact basement membrane for tissue-specific gene expression. Sympson, C.J., Talhouk, R.S., Alexander, C.M., Chin, J.R., Clift, S.M., Bissell, M.J., Werb, Z. J. Cell Biol. (1994) [Pubmed]
  27. Differential interactions of specific nuclear factor I isoforms with the glucocorticoid receptor and STAT5 in the cooperative regulation of WAP gene transcription. Mukhopadhyay, S.S., Wyszomierski, S.L., Gronostajski, R.M., Rosen, J.M. Mol. Cell. Biol. (2001) [Pubmed]
  28. Production of active bovine tracheal antimicrobial peptide in milk of transgenic mice. Yarus, S., Rosen, J.M., Cole, A.M., Diamond, G. Proc. Natl. Acad. Sci. U.S.A. (1996) [Pubmed]
  29. Mammary gland specific hEGF receptor transgene expression induces neoplasia and inhibits differentiation. Brandt, R., Eisenbrandt, R., Leenders, F., Zschiesche, W., Binas, B., Juergensen, C., Theuring, F. Oncogene (2000) [Pubmed]
  30. Ectopic TGF beta 1 expression in the secretory mammary epithelium induces early senescence of the epithelial stem cell population. Kordon, E.C., McKnight, R.A., Jhappan, C., Hennighausen, L., Merlino, G., Smith, G.H. Dev. Biol. (1995) [Pubmed]
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