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Gene Review

cad  -  caudal

Drosophila melanogaster

Synonyms: 38E.19, CAD, CG1759, Cad, Dmel\CG1759, ...
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Disease relevance of cad

  • CDX2, a human homologue of Drosophila caudal, is mutated in both alleles in a replication error positive colorectal cancer [1].
  • Cells of the rostral and intermediate vesicle contribute to the frontal ganglion; the hypocerebral ganglion develops from the intermediate vesicle, the paraesophageal ganglion from the rostral vesicle, and the ventricular ganglion from the caudal vesicle [2].

High impact information on cad

  • Translational inhibition depends on the Bcd binding region (BBR) present in the cad 3' untranslated region [3].
  • Thus, simultaneous interactions of d4EHP with the cap structure and of Bcd with BBR renders cad mRNA translationally inactive [3].
  • In third instar larvae, cad is expressed in the gut, the gonads, and parts of the genital discs [4].
  • Expression of the caudal gene in the germ line of Drosophila: formation of an RNA and protein gradient during early embryogenesis [4].
  • In the absence of the Hh pathway, cad induces internal analia development, probably by activating the brachyenteron and even-skipped genes [5].

Biological context of cad

  • This suggests that bcd may act in the region-specific control of cad mRNA translation [6].
  • Maternally-derived cad protein in the anterior region of embryos interferes with head morphogenesis, showing that cad mRNA suppression by Bcd is an important control event during early Drosophila embryogenesis [7].
  • Rather, we find that binding sites for other activators, including Hunchback and Caudal correlate with CRM expression in middle and posterior body regions [8].
  • The effects of heat-shock-induced ectopic expression of the homeobox gene caudal (cad) at all stages of Drosophila development have been examined [9].
  • Reduction of Gb' cad expression level by RNA interference resulted in deletion of the gnathum, thorax, and abdomen in embryos, remaining only anterior head [10].

Anatomical context of cad

  • In the absence of cad function, the analia develop like the immediately anterior segment (male genitalia), following the transformation rule of the canonical Hox genes [5].
  • The principal features of wingless expression patterns in Drosophila are conserved in Tribolium, including its early deployment in rostral and caudal domains in the blastoderm, its segmental iteration in cells immediately anterior to cells expressing the engrailed gene, and its later restriction to a ventral sector of the developing appendages [11].
  • Xpbx1b expression overlaps with Xmeis1 in several areas, including the lateral neural folds, caudal branchial arch, hindbrain, and optic cup [12].
  • Immediately subjacent to Sonic expression in the caudal endoderm is undifferentiated mesoderm, later to become the visceral mesoderm of the hindgut [13].
  • Role of caudal in hindgut specification and gastrulation suggests homology between Drosophila amnioproctodeal invagination and vertebrate blastopore [14].

Associations of cad with chemical compounds

  • Recently the ligands for Eph type receptor tyrosine kinases have been shown to be expressed strongly at the caudal tectum and play a role in retinotectal map formation by repulsing the temporal retinal fibers [15].

Physical interactions of cad

  • In this study, we found three potential Caudal binding sequences in the 5'-flanking region of the Drosophila E2F (DE2F) gene and showed by transient transfection assays that they are involved in Caudal transactivation of the dE2F gene promoter [16].
  • The effect of lethal mutations and deletions within the bithorax complex upon the identity of caudal metameres in the Drosophila embryo [17].

Enzymatic interactions of cad

  • The pathway of tissue specific gene regulation, apparently, branches beyond Krüppel to form at least a cut and a caudal branch [18].

Regulatory relationships of cad

  • We show here that the abdominal cad domain is regulated by the hunchback (hb) gradient through repression at high concentrations and activation at low concentrations of HB protein [19].
  • Employing this system, we show that the cad domain functions by activating the expression of the abdominal gap genes knirps (kni) and giant (gt) [19].
  • Caudal-induced E2F expression was also confirmed with a GAL4-UAS system in living flies [16].
  • These Abd-B-type Hox genes have been shown to be expressed in the most caudal regions of the developing vertebrate embryo and in overlapping domains within the developing limbs, suggesting that these genes play important roles in pattern formation in both appendicular and axial regions of the body [20].

Other interactions of cad

  • Here we show that caudal, a conserved homeodomain protein that forms a posterior-to-anterior concentration gradient, and the anterior determinant bicoid cooperate to form a partly redundant activator system in the posterior region of the embryo [21].
  • We show that translational repression of cad mRNA is dependent on a functional eIF4E-binding motif [22].
  • These results suggest that Caudal is involved in regulation of proliferation through transactivation of the E2F gene in Drosophila [16].
  • Furthermore, Gb' cad was found to be involved in the posterior elongation, acting as a downstream gene in the Wingless/Armadillo signalling pathways [10].
  • In addition, we found that the three primordia of the larval genital disc have already been specified in the GDPCs by the coordinated actions of the homeotic (Hox) genes, abdominal-A, Abdominal-B, and caudal [23].

Analytical, diagnostic and therapeutic context of cad


  1. CDX2, a human homologue of Drosophila caudal, is mutated in both alleles in a replication error positive colorectal cancer. Wicking, C., Simms, L.A., Evans, T., Walsh, M., Chawengsaksophak, K., Beck, F., Chenevix-Trench, G., Young, J., Jass, J., Leggett, B., Wainwright, B. Oncogene (1998) [Pubmed]
  2. Embryonic development of the stomatogastric nervous system in Drosophila. Hartenstein, V., Tepass, U., Gruszynski-Defeo, E. J. Comp. Neurol. (1994) [Pubmed]
  3. A new paradigm for translational control: inhibition via 5'-3' mRNA tethering by Bicoid and the eIF4E cognate 4EHP. Cho, P.F., Poulin, F., Cho-Park, Y.A., Cho-Park, I.B., Chicoine, J.D., Lasko, P., Sonenberg, N. Cell (2005) [Pubmed]
  4. Expression of the caudal gene in the germ line of Drosophila: formation of an RNA and protein gradient during early embryogenesis. Mlodzik, M., Gehring, W.J. Cell (1987) [Pubmed]
  5. Caudal is the Hox gene that specifies the most posterior Drosophile segment. Moreno, E., Morata, G. Nature (1999) [Pubmed]
  6. RNA binding and translational suppression by bicoid. Rivera-Pomar, R., Niessing, D., Schmidt-Ott, U., Gehring, W.J., Jäckle, H. Nature (1996) [Pubmed]
  7. Sequence interval within the PEST motif of Bicoid is important for translational repression of caudal mRNA in the anterior region of the Drosophila embryo. Niessing, D., Dostatni, N., Jäckle, H., Rivera-Pomar, R. EMBO J. (1999) [Pubmed]
  8. The role of binding site cluster strength in Bicoid-dependent patterning in Drosophila. Ochoa-Espinosa, A., Yucel, G., Kaplan, L., Pare, A., Pura, N., Oberstein, A., Papatsenko, D., Small, S. Proc. Natl. Acad. Sci. U.S.A. (2005) [Pubmed]
  9. Effects of ectopic expression of caudal during Drosophila development. Mlodzik, M., Gibson, G., Gehring, W.J. Development (1990) [Pubmed]
  10. caudal is required for gnathal and thoracic patterning and for posterior elongation in the intermediate-germband cricket Gryllus bimaculatus. Shinmyo, Y., Mito, T., Matsushita, T., Sarashina, I., Miyawaki, K., Ohuchi, H., Noji, S. Mech. Dev. (2005) [Pubmed]
  11. Conservation of wingless patterning functions in the short-germ embryos of Tribolium castaneum. Nagy, L.M., Carroll, S. Nature (1994) [Pubmed]
  12. Xpbx1b and Xmeis1b play a collaborative role in hindbrain and neural crest gene expression in Xenopus embryos. Maeda, R., Ishimura, A., Mood, K., Park, E.K., Buchberg, A.M., Daar, I.O. Proc. Natl. Acad. Sci. U.S.A. (2002) [Pubmed]
  13. Sonic hedgehog is an endodermal signal inducing Bmp-4 and Hox genes during induction and regionalization of the chick hindgut. Roberts, D.J., Johnson, R.L., Burke, A.C., Nelson, C.E., Morgan, B.A., Tabin, C. Development (1995) [Pubmed]
  14. Role of caudal in hindgut specification and gastrulation suggests homology between Drosophila amnioproctodeal invagination and vertebrate blastopore. Wu, L.H., Lengyel, J.A. Development (1998) [Pubmed]
  15. En-2 regulates the expression of the ligands for Eph type tyrosine kinases in chick embryonic tectum. Shigetani, Y., Funahashi, J.I., Nakamura, H. Neurosci. Res. (1997) [Pubmed]
  16. The caudal homeodomain protein activates Drosophila E2F gene expression. Hwang, M.S., Kim, Y.S., Choi, N.H., Park, J.H., Oh, E.J., Kwon, E.J., Yamaguchi, M., Yoo, M.A. Nucleic Acids Res. (2002) [Pubmed]
  17. The effect of lethal mutations and deletions within the bithorax complex upon the identity of caudal metameres in the Drosophila embryo. Whittle, J.R., Tiong, S.Y., Sunkel, C.E. Journal of embryology and experimental morphology. (1986) [Pubmed]
  18. Regulatory interactions and role in cell type specification of the Malpighian tubules by the cut, Krüppel, and caudal genes of Drosophila. Liu, S., Jack, J. Dev. Biol. (1992) [Pubmed]
  19. Zygotic caudal regulation by hunchback and its role in abdominal segment formation of the Drosophila embryo. Schulz, C., Tautz, D. Development (1995) [Pubmed]
  20. Homeotic transformations and limb defects in Hox A11 mutant mice. Small, K.M., Potter, S.S. Genes Dev. (1993) [Pubmed]
  21. Activation of posterior gap gene expression in the Drosophila blastoderm. Rivera-Pomar, R., Lu, X., Perrimon, N., Taubert, H., Jäckle, H. Nature (1995) [Pubmed]
  22. Bicoid associates with the 5'-cap-bound complex of caudal mRNA and represses translation. Niessing, D., Blanke, S., Jäckle, H. Genes Dev. (2002) [Pubmed]
  23. Allocation and specification of the genital disc precursor cells in Drosophila. Chen, E.H., Christiansen, A.E., Baker, B.S. Dev. Biol. (2005) [Pubmed]
  24. brachyenteron is necessary for morphogenesis of the posterior gut but not for anteroposterior axial elongation from the posterior growth zone in the intermediate-germband cricket Gryllus bimaculatus. Shinmyo, Y., Mito, T., Uda, T., Nakamura, T., Miyawaki, K., Ohuchi, H., Noji, S. Development (2006) [Pubmed]
  25. Transcriptional regulation of the Drosophila caudal homeobox gene by DRE/DREF. Choi, Y.J., Choi, T.Y., Yamaguchi, M., Matsukage, A., Kim, Y.S., Yoo, M.A. Nucleic Acids Res. (2004) [Pubmed]
  26. A maternal homeobox gene, Bombyx caudal, forms both mRNA and protein concentration gradients spanning anteroposterior axis during gastrulation. Xu, X., Xu, P.X., Suzuki, Y. Development (1994) [Pubmed]
  27. Scanning electron microscopy of Drosophila melanogaster embryogenesis. III. Formation of the head and caudal segments. Turner, F.R., Mahowald, A.P. Dev. Biol. (1979) [Pubmed]
  28. Expression of two even-skipped genes eve1 and evx2 during zebrafish fin morphogenesis and their regulation by retinoic acid. Brulfert, A., Monnot, M.J., Géraudie, J. Int. J. Dev. Biol. (1998) [Pubmed]
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