odd-paired

DEVELOPMENTAL BIOLOGY

Embryonic

See the embryonic expression pattern of opa at the Berkeley Drosophila Genome Project Patterns of Gene Expression Site.

opa transcript first appears at the beginning of cellularization (stage 5) in a single stripe located 80% of the distance to the anterior end of the embryo. With time its expression extends to form a large area reaching to 20% from the posterior end. Only at stage 7, once germ band expression begins, does the transcript begin to fade, generating 14 weak stripes over a low background level of expression (Benedyk, 1994).

As development proceeds, opa expression ceases briefly both in the ectoderm and in the underlying mesodermal cells that later become the visceral mesoderm (Cimbora, 1995).

Effects of Mutation or Deletion

Only half the number of denticle belts are present in opa mutants as a result of deletion of alternate segments (Benedyk, 1994).

odd-paired is required for normal midgut development. opa function is required for formation of the three characteristic midgut constrictions. In the cellular blastoderm, opa is expressed ubiquitously in the ectoderm and mesoderm precursors throughout the presumptive segmented region. As development proceeds, opa expression ceases briefly both in the ectoderm and in the underlying mesodermal cells that later become the visceral mesoderm. In opa mutants the visceral mesoderm is interrupted, evidently due to abnormal expression of bagpipe, a homeodomain gene required for the formation of the visceral mesoderm. At early stages of development in opa mutants, interruptions in the visceral mesoderm are observed at many positions along the anterior-posterior axis. As development proceeds, interruptions are less frequently observed; however, one interruption is coincident with the Antennapedia expression domain and variability of Ultrabithorax expression in opa mutants. From these observations, it is inferred that the loss of at least the first and second midgut constrictions in opa mutants is the result of defects first evident in the early stages of visceral mesoderm development (Cimbora, 1995).

During later stages in development, opa expression reinitiates in spatially restricted domains of the visceral mesoderm, coinciding with the locations at which the first and third constrictions will form. At the locations of the first and third constrictions, opa is positively regulated by Antennapedia and abdominal-A, respectively, while between these domains opa is negatively regulated by Ultrabithorax and dpp. The coincidence of opa expression with the locations of the first and third midgut constrictions and the complex regulation of opa expression leads to a speculation that in addition to contributing to visceral msoderm development, opa, acting in the mesoderm, may mediate homeotic gene function during midgut constriction formation (Cimbora, 1995).

In addition to its requirement for visceral mesoderm development, opa is required for proper gut expression in the endodermal component of the midgut. Repression of the POU-domain gene pdm-1 in a specific region of the endoderm is dependent on opa function. pdm-1 is expressed in much of the endoderm at stage 13, but is repressed in a domain of the endoderm underlying the Ubx and dpp expression domain in the visceral mesoderm. Repression of pdm-1 in the endoderm is dependent of Ubx and dpp, and the accumulation of DPP protein in this region of the endoderm suggests that it may be the secreted factor responsible for regulating pdm-1 across germ layers. In opa mutants, pdm-1 repression does not occur. opa does not appear to contribute to pdm-1 repression through Ubx or dpp, as these genes are expressed in opa mutants, and the activation of the homeotic gene labial in the endoderm (which is dependent on Ubx/dpp signaling) occurs normally. These observations suggest that repression of pdm-1 occurs by a novel signaling pathway, independent of dpp. Since opa is not transcribed in the visceral mesoderm near the pdm-1 expression domain, interruptions in the endoderm may reflect the inability of the endodermal primordia to migrate properly along the defective visceral mesoderm, an interpretation consistent with the proposal that the visceral mesoderm is required as a substrate for endodermal migration (Affolter, 1993 and Cimbora, 1995).

Affolter, M., Walldorf, U., Kloter, U., Schier, A.F. and Gehring, W.J. (1993). Regional repression of a Drosophila POU box gene in the endoderm involves inductive interactions between germ layers. Development 117: 1199-1210

Alper, S. and Kenyon, C. (2002). The zinc finger protein REF-2 functions with the Hox genes to inhibit cell fusion in the ventral epidermis of C. elegans. Development 129: 3335-3348. 12091304

Aruga, J., et al. (1996a). The mouse zic gene family. Homologues of the Drosophila pair-rule gene odd-paired. J. Biol. Chem. 271: 1043-7

Aruga, J., et al. (1996b). Identification and characterization of Zic4, a new member of the mouse Zic gene family. Gene 172: 291-294

Aruga, J., et al. (1998). Mouse zic1 is involved in cerebellar development. J. Neurosci. 18(1): 284-293.

Aruga, J., et al. (1999). Zic1 regulates the patterning of vertebral arches in cooperation with Gli3. Mech. of Dev. 89: 141-150.

Aruga, J., Shimoda, K. and Mikoshiba, K. (2000). A 5' segment of the mouse Zic1 gene contains a region specific enhancer for dorsal hindbrain and spinal cord. Brain Res. Mol. Brain Res. 78(1-2): 15-25. 10891581

Aruga, J., et al. (2002). Zic2 controls cerebellar development in cooperation with Zic1. J. Neurosci. 22(1): 218-225. 11756505

Aruga, J., et al. (2002). Zic1 promotes the expansion of dorsal neural progenitors in spinal cord by inhibiting neuronal differentiation. Dev. Biol. 244: 329-341. 11944941

Bagutti, C., et al. (2003). The intracellular domain of teneurin-2 has a nuclear function and represses zic-1-mediated transcription. J. Cell Sci. 116: 2957-2966. 12783990

Baumgartner, S.,Martin, D.,Hagios, C. and Chiquet-Ehrismann, R. (1994). Tenm, a Drosophila gene related to tenascin, is a new pair-rule gene. EMBO J 13: 3728-3740

Benedyk, M. J., Mullen, J. R. and DiNardo, S. (1994). odd-paired: a zinc finger pair-rule protein required for the timely activation of engrailed and wingless in Drosophila embryos. Genes Dev 8: 105-17

Bilder, D., Graba, Y. and Scott, M. (1998). Wnt and TGFbeta signals subdivide the AbdA hox domain during Drosophila mesoderm patterning. Development 125(9): 1781-1790.

Brown, S. A., et al. (1998). Holoprosencephaly due to mutations in ZIC2, a homologue of Drosophila odd-paired. Nat. Genet. 20(2): 180-3.

Cimbora, D. M. and Sakonju, S. (1995). Drosophila midgut morphogenesis requires the function of the segmentation gene odd-paired. Dev. Biol. 169: 580-595

de la Calle-Mustienes, E., et al. (2002). Xiro homeoproteins coordinate cell cycle exit and primary neuron formation by upregulating neuronal-fate repressors and downregulating the cell-cycle inhibitor XGadd45-gamma. Mech. Dev. 119: 69-80. 12385755

Ebert, P. J., et al. (2003). Zic1 represses Math1 expression via interactions with the Math1 enhancer and modulation of Math1 autoregulation. Development 130: 1949-1959. 12642498

Feledy, J. A., et al. (1999). Inhibitory patterning of the anterior neural plate in Xenopus by homeodomain factors Dlx3 and Msx1. Dev. Biol. 212(2): 455-64.

Franco, P. G., et al. (1999). Functional association of retinoic acid and hedgehog signaling in Xenopus primary neurogenesis. Development 126: 4257-4265.

Grienenberger, A., et al. (2003). Tgfß signaling acts on a Hox response element to confer specificity and diversity to Hox protein function. Development 130: 5445-5455. 14507783

Grinblat, Y., Gamse, J., Patel, M. and Sive, H. (1998). Determination of the zebrafish forebrain: induction and patterning. Development 125(22): 4403-4416

Imai, K. S., Satou, Y. and Satoh, N. (2002). Multiple functions of a Zic-like gene in the differentiation of notochord, central nervous system and muscle in Ciona savignyi embryos. Development 129: 2723-2732. 12015299

Kitaguchi, T., et al. (2000). Zic3 is involved in the left-right specification of the Xenopus embryo. Development 127: 4787-4795

Klingler, M. and Gergen, J. P. (1993). Regulation of runt transcription by Drosophila segmentation genes. Mech Dev 43: 3-19

Kobayashi, K., et al. (2003). Maternal macho-1 is an intrinsic factor that makes cell response to the same FGF signal differ between mesenchyme and notochord induction in ascidian embryos. Development 130: 5179-5190. 12954719

Koyabu, Y., et al. (2001). Physical and functional interactions between Zic and Gli proteins. J. Biol. Chem. 276(10): 6889-92. 11238441

Kuo, J. S., et al. (1998). opl: a zinc finger protein that regulates neural determination and patterning in Xenopus. Development 125(15): 2867-2882.

Leclerc, C., et al. (2003). Calcium transients triggered by planar signals induce the expression of ZIC3 gene during neural induction in Xenopus. Dev. Biol. 261: 381-390. 14499648

Lee, H., Stultz, B. G. and Hursh, D. A. (2007). The Zic family member, odd-paired, regulates the Drosophila BMP, decapentaplegic, during adult head development. Development 134(7): 1301-10. Medline abstract: 17329368

Lindgens, D., Holstein, T. W. and Technau, U. (2004). Hyzic, the Hydra homolog of the zic/odd-paired gene, is involved in the early specification of the sensory nematocytes. Development 131: 191-201. 14681184

Merzdorf, C. S. and Sive, H. L. (2006). The zic1 gene is an activator of Wnt signaling. Int. J. Dev. Biol. 50(7): 611-7. Medline abstract: 16892174

Mizugishi, K., et al. (2001). Molecular properties of Zic proteins as transcriptional regulators and their relationship to GLI proteins. J. Biol. Chem. 276(3): 2180-8. 11053430

Mizuseki, K., et al. (1998). Xenopus Zic-related-1 and Sox-2, two factors induced by chordin, have distinct activities in the initiation of neural induction. Development 125(4): 579-587.

Nagai, T., et al. (1997). The expression of the mouse Zic1, Zic2 and Zic3 genes suggests an essential role for Zic genes in body pattern formation. Dev. Biol. 182: 299-313

Nagai, T., et al. (2000). Zic2 regulates the kinetics of neurulation. Proc. Natl. Acad. Sci. 97: 1618-1623.

Nakata, K., et al. (1997). Xenopus zic3, a primary regulator both in neural and neural crest development. Proc. Natl. Acad. Sci. 94(22): 11980-11985.

Nakata, K., et al. (2000). A novel member of the Xenopus Zic family, Zic5, mediates neural crest development. Mech. Dev. 99: 83-91.

Nyholm, M. K., Wu, S. F., Dorsky, R. I. and Grinblat, Y. (2007). The zebrafish zic2a-zic5 gene pair acts downstream of canonical Wnt signaling to control cell proliferation in the developing tectum. Development 134(4): 735-46. Medline abstract: 17215296

Pak, W., Hindges, R., Lim, Y. S., Pfaff, S. L. and O'Leary, D. D. (2004). Magnitude of binocular vision controlled by islet-2 repression of a genetic program that specifies laterality of retinal axon pathfinding. Cell 119(4): 567-78. 15537545

Rohr, K. B., Schulte-Merker, S. and Tautz, D. (1999). Zebrafish zic1 expression in brain and somites is affected by BMP and Hedgehog signalling. Mec. Dev. 85(1-2): 147-159.

Salero, E., et al. (2001). Transcription factors Zic1 and Zic2 bind and transactivate the apolipoprotein E gene promoter. J. Biol. Chem. 276(3): 1881-8. 11038359

Sassa, T., Aizawa, H. and Okamoto, H. (2007). Visualization of two distinct classes of neurons by gad2 and zic1 promoter/enhancer elements in the dorsal hindbrain of developing zebrafish reveals neuronal connectivity related to the auditory and lateral line systems. Dev. Dyn. 236(3): 706-18. Medline abstract: 17279576

Sato, T., Sasai, N. and Sasai, Y. (2005). Neural crest determination by co-activation of Pax3 and Zic1 genes in Xenopus ectoderm. Development 132: 2355-2363. 15843410

Skeath, J. B., et al. (1992). Gene regulation in two dimensions: the proneural achaete-scute genes are controlled by combinations of axis-patterning genes through a common intergenic control region. Genes and Dev. 6: 2606-19

Swantek, D. and Gergen, J. P. (2004). Ftz modulates Runt-dependent activation and repression of segment-polarity gene transcription. Development 131: 2281-229015102703

Ursuliak, Z., et al (1997). Differential accumulation of DPTP61F alternative transcripts: regulation of a protein tyrosine phosphatase by segmentation genes. Mech. Dev. 65(1-2): 19-30.

Wada, S. and Saiga, H. (2002). HrzicN, a new Zic family gene of ascidians, plays essential roles in the neural tube and notochord development. Development 129: 5597-5608. 12421701

Yagi, K., Satou, Y. and Satoh, N. (2004). A zinc finger transcription factor, ZicL, is a direct activator of Brachyury in the notochord specification of Ciona intestinalis. Development 131: 1279-1288. 14993185

Zhang, J., Jin, Z and Bao, Z.-Z. (2004). Disruption of gradient expression of Zic3 resulted in abnormal intra-retinal axon projection. Development 131: 1553-1562. 14985256

date revised: 20 August 2007

Home page: The Interactive Fly © 1997 Thomas B. Brody, Ph.D.

The Interactive Fly resides on the
Society for Developmental Biology's Web server.