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Gene Review

mixl1  -  Mix paired-like homeobox

Danio rerio

Synonyms: SO:0000704, bon, bon/mixer, mix, mix-like
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High impact information on bon

  • In this report, we first show that the zebrafish bonnie and clyde (bon) gene plays a critical early role in endoderm formation. bon mutants exhibit a profound reduction in the number of sox17-expressing endodermal precursors formed during gastrulation, and, consequently, a profound reduction in gut tissue at later stages [1].
  • The endodermal precursors that do form in bon mutants, however, appear to differentiate normally indicating that bon is not required at later steps of endoderm development [1].
  • Overexpression of bon can rescue endodermal gene expression and the formation of a gut tube in bon mutants [1].
  • Consistent with these results, bon-mutant embryos show defects in axial mesoderm gene expression starting at mid-gastrulation stages [2].
  • We find that sur expression is independent of Nodal signaling and that bon is expressed in the absence of Nodal signaling but requires Nodal signaling and Sur for enhanced, maintained expression [3].

Biological context of bon

  • In contrast to the relatively mild defects observed in single mutants, loss of both bon and sur results in a severe phenotype characterized by absence of prechordal plate, cardiac mesoderm, endoderm and ventral neuroectoderm [3].
  • Smoothing of the enveloping cell layer at the midblastula transition occurred normally and expression of bon/mixer and mezzo, zygotic transcription factors indicated that activation of the zygotic genome did occur [4].

Anatomical context of bon

  • Here, we show that, in addition to its previously described role in endoderm formation, Bon also regulates the anteroposterior patterning of the neuroectoderm. bon-mutant embryos exhibit an anterior reduction of the neural plate [2].


  1. The zebrafish bonnie and clyde gene encodes a Mix family homeodomain protein that regulates the generation of endodermal precursors. Kikuchi, Y., Trinh, L.A., Reiter, J.F., Alexander, J., Yelon, D., Stainier, D.Y. Genes Dev. (2000) [Pubmed]
  2. The Mix family homeodomain gene bonnie and clyde functions with other components of the Nodal signaling pathway to regulate neural patterning in zebrafish. Trinh, L.A., Meyer, D., Stainier, D.Y. Development (2003) [Pubmed]
  3. Mixer/Bon and FoxH1/Sur have overlapping and divergent roles in Nodal signaling and mesendoderm induction. Kunwar, P.S., Zimmerman, S., Bennett, J.T., Chen, Y., Whitman, M., Schier, A.F. Development (2003) [Pubmed]
  4. Role of Fyn kinase in signaling associated with epiboly during zebrafish development. Sharma, D., Holets, L., Zhang, X., Kinsey, W.H. Dev. Biol. (2005) [Pubmed]
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