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Gene Review

Med  -  Medea

Drosophila melanogaster

Synonyms: CG1775, Dmel\CG1775, E(zen)3, MED, SMAD4, ...
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High impact information on Med

  • Furthermore, we present genetic and biochemical evidence that the brk silencer serves as a direct target for a protein complex consisting of the Smad homologs Mad/Medea and the zinc finger protein Schnurri [1].
  • Dpp signals via the SMAD proteins Mad and Medea [2].
  • Screens for binding factors yielded Tinman itself and the Smad4 homolog Medea [3].
  • Here we show that Hiw binds to the Smad protein Medea (Med) [4].
  • Indeed, ectopic expression of a dad-LacZ enhancer trap revealed that the Dpp pathway was upregulated in slmb somatic clones and, consistent with this, ectopic accumulation of the co-Smad protein, Medea, was recorded [5].

Biological context of Med


Anatomical context of Med

  • Furthermore, inactivating MEDEA mutations prevent nuclear translocation either by preventing interaction with MAD or by trapping MAD/MEDEA complexes in the cytosol [6].
  • Genetic analysis suggests that Medea may have two independently mutable functions in patterning the embryonic ectoderm [10].
  • Our data demonstrate that nuclear accumulation of the co-SMAD Medea requires a BMP signal during blastoderm and gastrula stages [11].
  • We propose that dSno functions as a switch in optic lobe development, shunting Medea from the Dpp pathway to the Activin pathway to ensure proper proliferation [12].
  • As part of the study of this species, two diploid cell lines (2n = 12) of the Med-fly were established in vitro on two different culture media [13].

Associations of Med with chemical compounds

  • Binding was also affected by alanine substitutions in Mad and Med at a subset of basic residues within and flanking helix 2, indicating a contribution to binding of the GRCGNC and GTCT sites [8].

Regulatory relationships of Med

  • Screens for dpp interactions with maternally expressed genes have identified loss of function mutations in Mothers against dpp and Medea [14].

Other interactions of Med

  • In the wing primordium, loss of Medea most severely affects regions receiving low DPP signal [6].
  • Furthermore, we show that dSMAD2 complexes with MEDEA only in the presence of ATR-I and PUNT. dSmad2 is expressed in the imaginal disks and in the outer proliferation centre of the larval brain, suggesting that it may have important proliferative and patterning roles during Drosophila development [15].
  • We show that a weak, homozygous-viable sog mutant is enhanced to lethality by reduction in the activities of the Smad family members Mad or Medea, and that the lethality is caused by defects in the molecular specification and subsequent cellular differentiation of the dorsal-most cell type, the amnioserosa [16].
  • In marked GSCs mutant for Med and punt, two essential Bmp signal transducers, bam transcription is also elevated [17].


  1. Conversion of an extracellular Dpp/BMP morphogen gradient into an inverse transcriptional gradient. Müller, B., Hartmann, B., Pyrowolakis, G., Affolter, M., Basler, K. Cell (2003) [Pubmed]
  2. The Drosophila gene brinker reveals a novel mechanism of Dpp target gene regulation. Jaźwińska, A., Kirov, N., Wieschaus, E., Roth, S., Rushlow, C. Cell (1999) [Pubmed]
  3. Smad proteins act in combination with synergistic and antagonistic regulators to target Dpp responses to the Drosophila mesoderm. Xu, X., Yin, Z., Hudson, J.B., Ferguson, E.L., Frasch, M. Genes Dev. (1998) [Pubmed]
  4. Highwire regulates presynaptic BMP signaling essential for synaptic growth. McCabe, B.D., Hom, S., Aberle, H., Fetter, R.D., Marques, G., Haerry, T.E., Wan, H., O'Connor, M.B., Goodman, C.S., Haghighi, A.P. Neuron (2004) [Pubmed]
  5. Multiple roles of the F-box protein Slimb in Drosophila egg chamber development. Muzzopappa, M., Wappner, P. Development (2005) [Pubmed]
  6. Medea is a Drosophila Smad4 homolog that is differentially required to potentiate DPP responses. Wisotzkey, R.G., Mehra, A., Sutherland, D.J., Dobens, L.L., Liu, X., Dohrmann, C., Attisano, L., Raftery, L.A. Development (1998) [Pubmed]
  7. Regulation of spalt expression in the Drosophila wing blade in response to the Decapentaplegic signaling pathway. Barrio, R., de Celis, J.F. Proc. Natl. Acad. Sci. U.S.A. (2004) [Pubmed]
  8. Dpp-responsive silencers are bound by a trimeric Mad-Medea complex. Gao, S., Steffen, J., Laughon, A. J. Biol. Chem. (2005) [Pubmed]
  9. Smad6 is a Smad1/5-induced smad inhibitor. Characterization of bone morphogenetic protein-responsive element in the mouse Smad6 promoter. Ishida, W., Hamamoto, T., Kusanagi, K., Yagi, K., Kawabata, M., Takehara, K., Sampath, T.K., Kato, M., Miyazono, K. J. Biol. Chem. (2000) [Pubmed]
  10. The Drosophila Medea gene is required downstream of dpp and encodes a functional homolog of human Smad4. Hudson, J.B., Podos, S.D., Keith, K., Simpson, S.L., Ferguson, E.L. Development (1998) [Pubmed]
  11. Stepwise formation of a SMAD activity gradient during dorsal-ventral patterning of the Drosophila embryo. Sutherland, D.J., Li, M., Liu, X.Q., Stefancsik, R., Raftery, L.A. Development (2003) [Pubmed]
  12. dSno Facilitates Baboon Signaling in the Drosophila Brain by Switching the Affinity of Medea Away From Mad and Toward dSmad2. Takaesu, N.T., Hyman-Walsh, C., Ye, Y., Wisotzkey, R.G., Stinchfield, M.J., O'connor, M.B., Wotton, D., Newfeld, S.J. Genetics (2006) [Pubmed]
  13. Establishment of continuously in vitro growing cell lines of med-fly (Ceratitis capitata wied.). Cavalloro, R. Revue canadienne de biologie / éditée par l'Université de Montréal. (1981) [Pubmed]
  14. Genetic screens to identify elements of the decapentaplegic signaling pathway in Drosophila. Raftery, L.A., Twombly, V., Wharton, K., Gelbart, W.M. Genetics (1995) [Pubmed]
  15. Drosophila dSmad2 and Atr-I transmit activin/TGFbeta signals. Das, P., Inoue, H., Baker, J.C., Beppu, H., Kawabata, M., Harland, R.M., Miyazono, K., Padgett, R.W. Genes Cells (1999) [Pubmed]
  16. A positive role for Short gastrulation in modulating BMP signaling during dorsoventral patterning in the Drosophila embryo. Decotto, E., Ferguson, E.L. Development (2001) [Pubmed]
  17. Bmp signals from niche cells directly repress transcription of a differentiation-promoting gene, bag of marbles, in germline stem cells in the Drosophila ovary. Song, X., Wong, M.D., Kawase, E., Xi, R., Ding, B.C., McCarthy, J.J., Xie, T. Development (2004) [Pubmed]
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