GSC is first detected at cellularization (Stage 4) as a dorsoanterior stripe 2-3 cells wide, appearing in a strong horseshoe-like pattern across the dorsal side of the embryo anterior to the appearing cephalic furrow [Images]. This region corresponds to an extreme anterior-dorsal position of most insects, because the head neuraxis of Drosophila is bent dorsally. At germband extension (Stage 8) GSC is down-regulated in the dorsal-most region of the embryo and the horseshoe pattern resolves into a large cluster of cells on each side of the presumptive head. These positive cells migrate laterally and, by stage 10, GSC is restricted to three small clusters that migrate inside the embryo at germband retraction and are then associated with the brain hemispheres (Goriely, 1996 and Hahn, 1996).

A second domain of expression is seen as a cluster of cells at the time the stomodeum is first visible. This cluster follows the movements of foregut invagination and is incorporated into the roof of the stomodeum abutting the endodermic anterior midgut territory. These cells appear to be part of the anterior foregut, the ring gland and the stomatogastric nervous system, a tissue that is modified in a mutant lacking Gsc function (Goriely, 1996 and Hahn, 1996)


Abe, N., Dror, I., Yang, L., Slattery, M., Zhou, T., Bussemaker, H. J., Rohs, R. and Mann, R. S. (2015). Deconvolving the recognition of DNA shape from sequence. Cell 161(2): 307-318. PubMed ID: 25843630

Angerer, L. M., et al. (2001). Sea urchin goosecoid function links fate specification along the animal-vegetal and oral-aboral embryonic axes. Development 128: 4393-4404. 11714666

Armes, N. A. and Smith, J. C. (1997). The ALK-2 and ALK-4 activin receptors transduce distinct mesoderm-inducing signals during early Xenopus development but do not co-operate to establish thresholds. Development 124(19): 3797-3804. PubMed ID: 9367435

Artinger, M., et al. (1997). Interaction of goosecoid and brachyury in Xenopus mesoderm patterning. Mech. Dev. 65(1-2): 187-196. PubMed ID: 9256355

Belo, J. A., et al. (1998). The prechordal midline of the chondrocranium is defective in Goosecoid-1 mouse mutants. Mech. Dev. 72(1-2): 15-25. PubMed ID: 9533949

Broun, M., Sokol, S. and Bode, H. R. (1999). Cngsc, a homologue of goosecoid, participates in the patterning of the head, and is expressed in the organizer region of Hydra. Development 126: 5245-5254. PubMed ID: 10556050

Bruce, A. E. E., et al. (2003). The maternally expressed zebrafish T-box gene eomesodermin regulates organizer formation. Development 130: 5503-5517. 14530296

Candia, A. F. and Wright, C. V. (1995). The expression pattern of Xenopus Mox-2 implies a role in initial mesodermal differentiation. Mech Dev 52: 27-36. PubMed ID: 7577672

Conway, S. J. (1999). Novel expression of the Goosecoid transcription factor in the embryonic mouse heart. Mech. Dev. 81(1-2): 187-91. PubMed ID: 10330498

Croce, J., Lhomond, G. and Gache, C. (2003). Coquillette, a sea urchin T-box gene of the Tbx2 subfamily, is expressed asymmetrically along the oral-aboral axis of the embryo and is involved in skeletogenesis. Mech. Dev. 120: 561-572. 12782273

Ding, X, Hausen, P. and Steinbeisser, H. (1998). Pre-MBT patterning of early gene regulation in Xenopus: the role of the cortical rotation and mesoderm induction. Mech. Dev. 70: 15-24

Dixon, Fox, M. and Bruce, A. E. (2009). Short- and long-range functions of Goosecoid in zebrafish axis formation are independent of Chordin, Noggin 1 and Follistatin-like 1b. Development 136(10): 1675-85. PubMed Citation: 19369398

Ecochard, V., et al. (1998). A novel xenopus mix-like gene milk involved in the control of the endomesodermal fates. Development 125(14): 2577-2585. PubMed ID: 9510021

Fan, M. J. and Sokol, S. Y. (1997). A role for Siamois in Spemann organizer formation. Development 124(13): 2581-2589. PubMed ID: 9217000

Ferreiro, B., et al. (1998). Antimorphic goosecoids. Development 125(8): 1347-1359. PubMed Citation: 9502717

Filosa, S., et al. (1997). Goosecoid and HNF-3ß genetically interact to regulate neural tube patterning during mouse embryogenesis. Development 124(14): 2843-2854. PubMed Citation: 9226455

Galili, N., et al. (1998). Gscl, a gene within the minimal DiGeorge critical region, is expressed in primordial germ cells and the developing pons. Dev. Dyn. 212(1): 86-93. PubMed Citation: 9603426

Germain, S., et al. (2000). Homeodomain and winged-helix transcription factors recruit activated Smads to distinct promoter elements via a common Smad interaction motif. Genes Dev. 14: 435-451. PubMed Citation: 10691736

González-Gaitan, M. and Jäckle, H. (1995). Invagination centers within the Drosophila stomatogastric nervous system anlage are positioned by Notch-mediated signaling which is spatially controlled through wingless. Development 121: 2313-25. PubMed Citation: 7671798

Goriely, A., et al. (1996). A functional homologue of goosecoid in Drosophila. Development 122: 1641-1650. PubMed Citation: 8625850

Hahn, M. and Jackle, H. (1996). Drosophila goosecoid participates in neural development but not in body axis formation. EMBO J 15 (12): 3077-3084. PubMed Citation: 8670808

Hashimoto-Partyka, M. K., et al. (2003). Nodal signaling in Xenopus gastrulae is cell-autonomous and patterned by ß-Catenin. Dev. Bio. 253: 125-138. 12490202

Heanue, T. A., et al. (1997). Goosecoid misexpression alters the morphology and Hox gene expression of the developing chick limb bud. Mech. Dev. 69(1-2): 31-37

Jimenez, G., Verrijzer, C. P. and Ish-Horowicz, D. (1999). A conserved motif in Goosecoid mediates Groucho-dependent repression in Drosophila embryos. Mol. Cell. Biol. 19(3): 2080-7. PubMed Citation: 10022895

Jones, C. M., et al. (1995). Nodal-related signals induce axial mesoderm and dorsalize mesoderm during gastrulation. Development 121: 3651-62

Joore, J., et al (1997). Domains of retinoid signalling and neurectodermal expression of zebrafish otx1 and goosecoid are mutually exclusive. Biochem. Cell. Biol. 75(5): 601-612

Joore, J., et al. (1998). Protein kinase A is involved in the induction of early mesodermal marker genes by activin. Mech. Dev. 79(1-2): 5-16

Jung, C., Schnepf, M., Bandilla, P., Unnerstall, U. and Gaul, U. (2019). High sensitivity measurement of transcription factor-DNA binding affinities by competitive titration using fluorescence microscopy. J Vis Exp(144). PubMed ID: 30799844

Jung, C., Bandilla, P., von Reutern, M., Schnepf, M., Rieder, S., Unnerstall, U. and Gaul, U. (2018). True equilibrium measurement of transcription factor-DNA binding affinities using automated polarization microscopy. Nat Commun 9(1): 1605. PubMed ID: 29686282

Kessler, D. S. (1997). Siamois is required for formation of Spemann's organizer. Proc. Natl. Acad. Sci. 94(24): 13017-13022

Knoetgen, H., Viebahn, C. and Kessel, M. (1999). Head induction in the chick by primitive endoderm of mammalian, but not avian origin. Development 126(4): 815-825 >

Ku, M., et al. (2005). Positive and negative regulation of the transforming growth factor beta/activin target gene goosecoid by the TFII-I family of transcription factors. Mol. Cell. Biol. 25(16): 7144-57. 16055724

Labbe, E., et al. (1998). Smad2 and Smad3 positively and negatively regulate TGF beta-dependent transcription through the forkhead DNA-binding protein FAST2. Mol. Cell (1): 109-20

Lanctot, C., Lamolet, B. and Drouin, J. (1997). The bicoid-related homeoprotein Ptx1 defines the most anterior domain of the embryo and differentiates posterior from anterior lateral mesoderm. Development 124(14): 2807-2817

Latinkic, B. V. and Smith, J. C. (1999). Goosecoid and Mix.1 repress Brachyury expression and are required for head formation in Xenopus. Development 126(8): 1769-1779

Laurent, M. N., et al. (1997). The Xenopus homeobox gene twin mediates Wnt induction of goosecoid in establishment of Spemann's organizer. Development 124(23): 4905-4916

Lee, S. Y., Yoon, J., Lee, H. S., Hwang, Y. S., Cha, S. W., Jeong, C. H., Kim, J. I., Park, J. B., Lee, J. Y., Kim, S., Park, M. J., Dong, Z. and Kim, J. (2011). The function of heterodimeric AP-1 comprised of c-Jun and c-Fos in activin mediated Spemann organizer gene expression. PLoS One 6: e21796. PubMed ID: 21829441

Lemaire, L., et al. (1997). Segregating expression domains of two goosecoid genes during the transition from gastrulation to neurulation in chick embryos. Development 124: 1443-52

Li, J., Sagendorf, J. M., Chiu, T. P., Pasi, M., Perez, A. and Rohs, R. (2017). Expanding the repertoire of DNA shape features for genome-scale studies of transcription factor binding. Nucleic Acids Res 45(22): 12877-12887. PubMed ID: 29165643

Mailhos, C., Andre, S., Mollereau, B., Goriely, A., Hemmati-Brivanlou, A. and Desplan, C. (1998). Drosophila Goosecoid requires a conserved heptapeptide for repression of paired-class homeoprotein activators. Development 125: 937-947. PubMed ID: 9449676

McKendry, R., Harland, R. M. and Stachel, S. E. (1998). Activin-induced factors maintain goosecoid transcription through a paired homeodomain binding site. Dev. Biol. 204(1): 172-86.

Messenger, N. J., et al. (2005). Functional specificity of the Xenopus T-domain protein Brachyury is conferred by its ability to interact with Smad1. Dev. Cell 8(4): 599-610. 15809041

Mochizuki, T., et al. (2000). Xlim-1 and LIM domain binding protein 1 cooperate with various transcription factors in the regulation of the goosecoid promoter. Dev. Bio. 224: 470-485. PubMed ID: 10926781

Niehrs, C., et al. (1993). The homeobox gene goosecoid controls cell migration in Xenopus embryos. Cell 72: 491-503

Papin, C. and Smith, J. C. (2000). Gradual refinement of activin-induced thresholds requires protein synthesis. Dev. Biol. 217: 166-172. PubMed ID: 10625543

Parry, D. A., Logan, C. V., Stegmann, A. P., Abdelhamed, Z. A., Calder, A., Khan, S., Bonthron, D. T., Clowes, V., Sheridan, E., Ghali, N., Chudley, A. E., Dobbie, A., Stumpel, C. T. and Johnson, C. A. (2013). SAMS, a syndrome of short stature, auditory-canal atresia, mandibular hypoplasia, and skeletal abnormalities is a unique neurocristopathy caused by mutations in Goosecoid. Am J Hum Genet. 93(6): 1135-42. PubMed ID: 24290375

Re'em-Kalma, Y., Lamb, T. and Frank, D. (1995). Competition between noggin and bone morphogenetic protein 4 activities may regulate dorsalization during Xenopus development. Proc. Natl. Acad. Sci. 92: 12141-12145

Reid, C. D., Zhang, Y., Sheets, M. D. and Kessler, D. S. (2012). Transcriptional integration of Wnt and Nodal pathways in establishment of the Spemann organizer. Dev Biol 368: 231-241. PubMed ID: 22627292

Ring, C., et al. (2002). The role of a Williams-Beuren syndrome-associated helix-loop-helix domain-containing transcription factor in activin/nodal signaling. Genes Dev. 16: 820-835. 11937490

Rivera-Perez, J. A., Wakamiya, M. and Behringer, R. R. (1999). Goosecoid acts cell autonomously in mesenchyme-derived tissues during craniofacial development. Development 126: 3811-3821. PubMed ID: 10433910

Rube, H. T., Rastogi, C., Kribelbauer, J. F. and Bussemaker, H. J. (2018). A unified approach for quantifying and interpreting DNA shape readout by transcription factors. Mol Syst Biol 14(2): e7902. PubMed ID: 29472273

Schmidt-Ott, et al. (1994). Number, identity, and sequence of the Drosophila head segments as revealed by neural elements and their deletion patterns in mutants. Proc. Natl. Acad. Sci. 91: 8363-8367. PubMed Citation: 7915837

Schulte-Merker, S., et al. (1994). Expression of zebrafish goosecoid and no tail gene products in wild-type and mutant no tail embryos. Development 120: 843-852

Smith, S. T. and Jaynes, J. B. (1996). A conserved region of Engrailed, shared among all en-, gsc-, Nk1-, Nk2- and msh-class homeoproteins, mediates active transcriptional repression in vivo. Development 122(10): 3141-3150

Sudou, N., Yamamoto, S., Ogino, H. and Taira, M. (2012). Dynamic in vivo binding of transcription factors to cis-regulatory modules of cer and gsc in the stepwise formation of the Spemann-Mangold organizer. Development 139: 1651-1661. PubMed ID: 22492356

Thisse, C., et al. (1994).Goosecoid expression in neurectoderm and mesendoderm is disrupted in zebrafish cyclops gastrulas. Dev Biol 164: 420-429

Tolkunova, E. N., Fujioka, M., Kobayashi, M., Deka, D. and Jaynes, J. B. (1998). Two distinct types of repression domain in engrailed: one interacts with the groucho corepressor and is preferentially active on integrated target genes. Mol Cell Biol 18: 2804-2814. PubMed ID: 9566899

Toyama, R., et al. (1995). Nodal induces ectopic goosecoid and lim1 expression and axis duplication in zebrafish. Development 121: 383-391

Tucker, A. S., et al. (2004). Bapx1 regulates patterning in the middle ear: altered regulatory role in the transition from the proximal jaw during vertebrate evolution. Development 131: 1235-1245. 14973294

Wacker, S., et al. (1998). Patterns and control of cell motility in the Xenopus gastrula. Development 125(10): 1931-1942. PubMed ID: 9550725

Yamada, G., et al. (1995). Targeted mutation of the murine goosecoid gene results in craniofacial defects and neonatal death. Development 121: 2917-2922

Yamanaka, Y., et al. (1998). A novel homeobox gene, dharma, can induce the organizer in a non-cell-autonomous manner. Genes Dev. 12(15): 2345-2353

Yao, J. and Kessler, D. S. (2001). Goosecoid promotes head organizer activity by direct repression of Xwnt8 in Spemann's organizer. Development 128: 2975-2987. 11532920

Yasuo, H. and Lemaire, P. (2001). Role of Goosecoid, Xnot and Wnt antagonists in the maintenance of the notochord genetic program in Xenopus gastrulae. Development 128: 3783-3793. 11585804

Zhou, T., Yang, L., Lu, Y., Dror, I., Dantas Machado, A. C., Ghane, T., Di Felice, R. and Rohs, R. (2013). DNAshape: a method for the high-throughput prediction of DNA structural features on a genomic scale. Nucleic Acids Res 41(Web Server issue): W56-62. PubMed ID: 23703209

Zhu, L., et al. (1999). Goosecoid regulates the neural inducing strength of the mouse node. Dev. Biol. 216(1): 276-81. PubMed ID: 10588878

Zoltewicz, J. S. and Gerhart, J. C. (1997). The Spemann organizer of Xenopus is patterned along its anteroposterior axis at the earliest gastrula stage. Dev. Biol. 192(2): 482-491

Goosecoid: Biological Overview | Evolutionary Homologs | Regulation | Developmental Biology

date revised: 20 April 2021 

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