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Gene Review

mixl1  -  Mix paired-like homeobox

Xenopus laevis

Synonyms: mix-a, mix.1, mix.2, mix1, mix1-a, ...
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High impact information on Mix.1


Biological context of Mix.1

  • We report the cloning, sequence analysis, and developmental expression pattern of lim1, a member of the LIM class homeobox gene family in the mouse. lim1 cDNA encodes a predicted 406 amino acid protein that is 93% identical with the product of the Xenopus LIM class homeobox gene Xlim1 [3].
  • Like the chromosomal Mix.1 gene, microinjected Mix.2 gene plasmids respond to activin in the presence of cycloheximide in animal cap assays and also respond to the embryonic inductive signal in Nieuwkoop recombinants [4].
  • We have determined the DNA sequence of a Xhox-1 cDNA and this allows us to predict the complete sequence of a vertebrate homeobox protein [5].
  • The sequence homology, expression pattern and gain-of-function phenotype of Xtwn is most similar to the previously isolated Xenopus homeobox gene siamois (Xsia) suggesting that Xtwn and Xsia comprise a new subclass of homeobox genes important in dorsal axis specification [6].
  • In Xenopus, this territory is initially included within the expression domain of the bicoid-class homeobox gene Xotx2 but very soon, at the beginning of neurulation, it becomes devoid of Xotx2 transcripts in spatiotemporal concomitance with the transcriptional activation of the paired-like homeobox gene Xrx1 [7].

Anatomical context of Mix.1

  • Next, injection of dsRNA for Xlim-1, a homeobox gene suggested to be involved in Spemann organizer functions, reduced the endogenous level of Xlim-1 mRNA and produced embryos with reduced eyes or anterior truncation at high efficiency [8].
  • This activity is necessary for normal expression of Mix.1, a mesoendodermal marker, in the endoderm as well as in the mesoderm, indicating that MAP kinase plays a functional role in patterning of both of these germ layers [9].
  • We generated domain swap mutants between Mix.3/Mixer and Mix.1 and tested their ability to induce endoderm in explanted ectoderm [10].
  • Expression of Xenopus Mix.3/Mixer in explanted ectoderm results in endoderm differentiation, whereas Mix.1 expression does not [10].
  • Interference with function of a homeobox gene in Xenopus embryos produces malformations of the anterior spinal cord [11].

Associations of Mix.1 with chemical compounds

  • The regional expression of two cell-specific markers, the homeobox protein Xhox3 and the neurotransmitter serotonin was also examined in embryos exposed to RA [12].
  • Under these conditions, expression of the anterior neural-specific homeobox gene engrailed is not detected, while the notochord-specific epitope recognized by the Tor-70 antibody is expressed in the presence of H2DIDS [13].

Other interactions of Mix.1

  • Xlim-1, a LIM class homeobox gene expressed in Xenopus laevis, is one of the earliest known marker genes of pronephros development and is expressed in pronephros rudiment [14].
  • This mutual repression is important for the specification of the embryonic body plan as ectopic expression of Mix.1 in the Xbra domain suppresses mesoderm differentiation [15].
  • We show that p53 functionally and physically interacts with the activin and bone morphogenetic protein pathways to directly induce the expression of the homeobox genes Xhox3 and Mix.1/2 [16].
  • However, activin-type signaling is present in vegetal cells, since the vegetal expression of Mix.1 and goosecoid is inhibited by the truncated activin receptor [17].
  • Expression of LIM class homeobox gene Xlim-3 in Xenopus development is limited to neural and neuroendocrine tissues [18].

Analytical, diagnostic and therapeutic context of Mix.1


  1. The homeobox gene goosecoid controls cell migration in Xenopus embryos. Niehrs, C., Keller, R., Cho, K.W., De Robertis, E.M. Cell (1993) [Pubmed]
  2. Mix.1, a homeobox mRNA inducible by mesoderm inducers, is expressed mostly in the presumptive endodermal cells of Xenopus embryos. Rosa, F.M. Cell (1989) [Pubmed]
  3. Expression patterns of the murine LIM class homeobox gene lim1 in the developing brain and excretory system. Fujii, T., Pichel, J.G., Taira, M., Toyama, R., Dawid, I.B., Westphal, H. Dev. Dyn. (1994) [Pubmed]
  4. DNA sequences mediating the transcriptional response of the Mix.2 homeobox gene to mesoderm induction. Vize, P.D. Dev. Biol. (1996) [Pubmed]
  5. Embryonic expression and nuclear localization of Xenopus homeobox (Xhox) gene products. Harvey, R.P., Tabin, C.J., Melton, D.A. EMBO J. (1986) [Pubmed]
  6. The Xenopus homeobox gene twin mediates Wnt induction of goosecoid in establishment of Spemann's organizer. Laurent, M.N., Blitz, I.L., Hashimoto, C., Rothbächer, U., Cho, K.W. Development (1997) [Pubmed]
  7. Role of Xrx1 in Xenopus eye and anterior brain development. Andreazzoli, M., Gestri, G., Angeloni, D., Menna, E., Barsacchi, G. Development (1999) [Pubmed]
  8. RNA interference for the organizer-specific gene Xlim-1 in Xenopus embryos. Nakano, H., Amemiya, S., Shiokawa, K., Taira, M. Biochem. Biophys. Res. Commun. (2000) [Pubmed]
  9. Localization of MAP kinase activity in early Xenopus embryos: implications for endogenous FGF signaling. LaBonne, C., Whitman, M. Dev. Biol. (1997) [Pubmed]
  10. Determination of the minimal domains of Mix.3/Mixer required for endoderm development. Doherty, J.R., Zhu, H., Kuliyev, E., Mead, P.E. Mech. Dev. (2006) [Pubmed]
  11. Interference with function of a homeobox gene in Xenopus embryos produces malformations of the anterior spinal cord. Wright, C.V., Cho, K.W., Hardwicke, J., Collins, R.H., De Robertis, E.M. Cell (1989) [Pubmed]
  12. Retinoic acid modifies the pattern of cell differentiation in the central nervous system of neurula stage Xenopus embryos. Ruiz i Altaba, A., Jessell, T.M. Development (1991) [Pubmed]
  13. An increase in intracellular pH during neural induction in Xenopus. Sater, A.K., Alderton, J.M., Steinhardt, R.A. Development (1994) [Pubmed]
  14. A role for Xlim-1 in pronephros development in Xenopus laevis. Chan, T.C., Takahashi, S., Asashima, M. Dev. Biol. (2000) [Pubmed]
  15. A role for the vegetally expressed Xenopus gene Mix.1 in endoderm formation and in the restriction of mesoderm to the marginal zone. Lemaire, P., Darras, S., Caillol, D., Kodjabachian, L. Development (1998) [Pubmed]
  16. Interplay between the tumor suppressor p53 and TGF beta signaling shapes embryonic body axes in Xenopus. Takebayashi-Suzuki, K., Funami, J., Tokumori, D., Saito, A., Watabe, T., Miyazono, K., Kanda, A., Suzuki, A. Development (2003) [Pubmed]
  17. FGF is a prospective competence factor for early activin-type signals in Xenopus mesoderm induction. Cornell, R.A., Musci, T.J., Kimelman, D. Development (1995) [Pubmed]
  18. Expression of LIM class homeobox gene Xlim-3 in Xenopus development is limited to neural and neuroendocrine tissues. Taira, M., Hayes, W.P., Otani, H., Dawid, I.B. Dev. Biol. (1993) [Pubmed]
  19. Molecular link in the sequential induction of the Spemann organizer: direct activation of the cerberus gene by Xlim-1, Xotx2, Mix.1, and Siamois, immediately downstream from Nodal and Wnt signaling. Yamamoto, S., Hikasa, H., Ono, H., Taira, M. Dev. Biol. (2003) [Pubmed]
  20. Microinjection of synthetic Xhox-1A homeobox mRNA disrupts somite formation in developing Xenopus embryos. Harvey, R.P., Melton, D.A. Cell (1988) [Pubmed]
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