During cell cycle 14, transcription of Trg commences throughout the posterior terminal region of the embryo. During cellularization, expression is down-regulated in the posterior tip, resulting in expression in a ring of cells that encompass the primordium of the hindgut and anal pads. Expression continues after invagination of the posterior midgut (Kispert, 1994 and Murakami, 1995).

Effects of Mutation or Deletion

After germ-band retraction in Trg deficient embryos, the major part of the hindgut is missing (Kispert, 1994 and Murakami, 1995).

Loss of Zn finger homeodomain 1 activity disrupts the development of two distinct mesodermal populations: the caudal visceral mesoderm (along which germ cells migrate) and the gonadal mesoderm (the final destination of the germ cells). The caudal visceral mesoderm facilitates the migration of germ cells from the endoderm to the mesoderm. Zfh-1 is also expressed in the gonadal mesoderm throughout the development of this tissue. Ectopic expression of Zfh-1 is sufficient to induce additional gonadal mesodermal cells and to alter the temporal course of gene expression within these cells. Germ cell migration was also analyzed in brachyenteron mutant embryos. Like zfh-1, byn is required for the migration of the caudal visceral mesoderm, but unlike zfh-1, it is not required for gonadal mesoderm development. Since byn and zfh-1 both disrupt caudal visceral mesoderm migration and show similar defects in germ cell migration, it is proposed that in wild-type embryos, the caudal visceral mesoderm facilitates the transition of many germ cells from the endoderm to the lateral mesoderm. abdominal-A is also required for gonadal mesoderm specification. Zfh-1 expression was analyzed in abdA mutants. Zfh-1 is expressed normally in mesodermal clusters at stage 10, however, its levels are not enhanced in PS10-12 during stage ll. The loss of high Zfh-1 expression correlates with the failure of SGP specification in abdA mutants. Although abdA is required for SGP specification, the initial stages of germ cell migration are unaffected in abdA mutant embryos (Broihier, 1998).

There are a number of mesodermal tissues that do not properly develop in embryos lacking the CVM, as in byn, fkh or tll embryos. For instance, the TVM develops aberrantly in byn mutants during late stages of embryogenesis. Although the inner layer of circular muscles differentiates in the absence of the CVM as in wild type, the morphogenesis of this layer does not proceed properly. The nuclei of the TVM are normally arranged as one broad band on each side of the midgut during germband retraction and subsequently split into two bands when the midgut primordia meet at stage 13. During this movement, the nuclei pass the rows of CVM cells, which are located at the dorsal and the ventral edge of the midgut primordium, respectively. In a byn mutant, however, the movement of the TVM nuclei is irregular, so that their organization into bands is lost and they become distributed over the entire gut circumference. Since byn is never expressed in the TVM, it is concluded that the proper arrangement and integrity of the circular muscle fibers requires the presence of the CVM. The irregular dorsoventral extension of the fibers results in an incomplete closure of the layer and the circular muscle layer of the midgut in byn embryos shows sporadic ruptures. These defects might be the reason why the three constrictions that normally subdivide the midgut tube into four gastric chambers are not formed in byn mutants. It seems rather unlikely that the longitudinal muscle fibers physically participate in the formation of the constrictions, since the fibers are oriented perpendicularly to the constriction planes (Kusch, 1999).

Strikingly, other mesodermal tissues that are affected in mutants lacking the CVM are not in obvious contact with the CVM during development. For instance, in byn mutants, the two rows of cardiac cells do not unite to form the heart vessel. In addition, pericardial cells are missing and the most dorsal internal muscle (dorsal acute 1: DA1) is absent or might be fused with DA2 in many segments. The progenitors of DA1 and of a subset of pericardial cells develop from a common cluster of dorsal mesodermal cells that can be followed from stage 10 on by their even-skipped (eve) expression. Three cells per hemisegment begin to express eve in each of 11 dorsal clusters in the mesoderm. By stage 12, the number of mesodermal eve cells increases by one in each cluster. This additional eve cell appears in succession from posterior to anterior clusters. Furthermore, it has been noted that the cells of the CVM pass the mesodermal eve clusters at a distance of about one cell diameter as they migrate anteriorly along the TVM. Shortly after the time when the leading edge of the CVM had passed, the fourth eve cell is added to the cluster. This addition occurs toward the CVM and by recruitment from neighboring cells rather than by cell division. Most importantly, the temporal and spatial correlation between the appearance of the fourth eve cell and the migration of the CVM is not a mere coincidence. In byn, tll or zfh-1 mutants in which the CVM fails to migrate anteriorly or is absent, the number of eve cells does not increase during germband retraction. It is proposed that this is the primary defect in the dorsal mesoderm that causes the defects in heart and dorsal muscle development of byn or tll mutants, and that normally an inductive signal emerging from the migrating CVM triggers the addition of the fourth eve cells. This view is supported by the observation that the specific rescue of CVM development in byn mutant embryos restores the dorsal mesodermal structures to a considerable extent. byn is neither expressed in the mesodermal eve cells nor in other dorsal mesodermal derivatives of the experimental embryos, but nevertheless the number and position of pericardial cells is essentially normal, the two rows of cardiac cells join and DA1 muscles are detectable in many segments (Kusch, 1999).

It was of interest to know whether byn is required solely for the early specification and migration of the CVM, or whether it is more directly involved in the signalling to the dorsal mesoderm. byn was therefore expressed outside the CVM, throughout the mesoderm, and the number of mesodermal eve cells was monitored. In such experimental embryos, a drastic increase of eve cells is seen at the dorsal edge of the mesoderm in the proximity to the original eve clusters during stage 11. Initially, these additional cells only appear close to the CVM, i.e. in the posterior half of the experimental embryos. Later, they also fill the gaps between the anterior eve clusters, to which the CVM fails to migrate upon ubiquitous mesodermal byn expression, and then form a band of cells along the entire dorsal mesoderm. Only the dorsal mesoderm appears to be competent to (directly or indirectly) respond to byn. This notion is supported by the finding that, in htl embryos that specifically lack derivatives of the dorsal mesoderm, ubiquitous mesodermal expression of byn does not lead to ectopic eve expression. Thus byn is not directly involved in transcriptionally activating eve in the dorsal mesoderm, since byn is normally never expressed in the eve clusters. Instead, it is proposed that byn regulates the expression of the ligand in the signalling process. byn can only exert this function on mesodermal cells, since a strictly ectodermal misexpression of byn has no effect on mesodermal eve expression. In fact, only cells in the neighborhood of the eve cells begin to express eve upon ubiquitous mesodermal byn expression, indicating that the competence to perceive the byn-mediated signal is dictated by contact with other eve cells (Kusch, 1999).


Amacher, S. L., et al. (2002). The zebrafish T-box genes no tail and spadetail are required for development of trunk and tail mesoderm and medial floor plate. Development 129: 3311-3323. 12091302

Amack, J. D. and Yost, H. J. (2004). The T box transcription factor no tail in ciliated cells controls zebrafish left-right asymmetry. Curr. Biol. 14: 685-690. 15084283

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 Citation: 9367435

Arnold, S. J., et al. (2000). Brachyury is a target gene of the Wnt/beta-catenin signaling pathway. Mech. Dev. 91: 249-258. PubMed Citation: 10704849

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

Bassham, S. and Postlethwait, J. (2000). Brachyury (T) expression in embryos of a larvacean urochordate, Oikopleura dioica, and the ancestral role of T. Dev. Biol. 220: 322-332. PubMed Citation: 10753519

Beck, C. W. and Slack, J. M. W. (1998). Analysis of the developing Xenopus tail bud reveals separate phases of gene expression during determination and growth. Mech. Dev. 72(1-2): 41-52. PubMed Citation: 9533951

Bielen, H., et al. (2007). Divergent functions of two ancient Hydra Brachyury paralogues suggest specific roles for their C-terminal domains in tissue fate induction. Development 134: 4187-4197. PubMed Citation: 17993466

Broihier, H. T., et al. (1998). zfh-1 is required for germ cell migration and gonadal mesoderm development in Drosophila. Development 125: 655-666. PubMed Citation: 9435286

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

Bulfone, A., et al. (1995). T-brain-1: a homolog of Brachyury whose expression defines molecularly distinct domains within the cerebral cortex. Neuron 15: 63-78. PubMed Citation: 7619531

Casey, E. S., et al. (1998). The T-box transcription factor Brachyury regulates expression of eFGF through binding to a non-palindromic response element. Development 125(19): 3887-3894. PubMed Citation: 9729496

Chiba, S., Jiang, D., Satoh, N. and Smith, W. C. (2009). Brachyury null mutant-induced defects in juvenile ascidian endodermal organs. Development 136(1):35-9. PubMed Citation: 19019990

Ciruna, B. and Rossant, J. (2001). FGF signaling regulates mesoderm cell fate specification and morphogenetic movement at the primitive streak. Dev. Cell 1: 37-49. PubMed Citation: 11703922

Conlon, F. L. and Smith, J. C. (1999). Interference with brachyury function inhibits convergent extension, causes apoptosis, and reveals separate requirements in the FGF and activin signalling pathways. Dev. Biol. 213(1): 85-100. PubMed Citation: 10452848

Corbo, J. C., Levine, M. and Zeller, R. W. (1997). Characterization of a notochord-specific enhancer from the Brachyury promoter region of the ascidian, Ciona intestinalis Development 124: 589-602. PubMed Citation: 9043074

Cunliffe, V. and Smith, J. C. (1994). Specification of mesodermal pattern in Xenopus laevis by interactions between Brachyury, noggin and Xwnt-8. EMBO J 13: 349-59. PubMed Citation: 7906224

Diaz, R. J., et al. (1996). Graded effect of tailless on posterior gut development: molecular basis of an allelic series of a nuclear receptor gene. Mech. Dev. 54: 119-130. PubMed Citation: 8808411

Di Gregorio, A. and Levine, M. (1999). Regulation of Ci-tropomyosin-like, a Brachyury target gene in the ascidian, Ciona intestinalis. Development 126: 5599-5609. PubMed Citation: 10572037

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. PubMed Citation: 9510021

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 Citation: 9636073

Erives, A. and Levine, M. (2000). Characterization of a maternal T-Box gene in Ciona intestinalis. Dev. Biol. 225: 169-178. PubMed Citation: 10964472

Kwan, K. M. and Kirschner, M. W. (2003). Xbra functions as a switch between cell migration and convergent extension in the Xenopus gastrula. Development 130: 1961-1972. 12642499

Fujiwara, S., Corbo, J. C. and Levine, M. (1998). The Snail repressor establishes a muscle/notochord boundary in the Ciona embryo. Development 125(13): 2511-2520. PubMed Citation: 9609834

Galceran, J., Hsu, S.-C. and Grosschedl, R. (2001). Rescue of a Wnt mutation by an activated form of LEF-1: Regulation of maintenance but not initiation of Brachyury expression. Proc. Natl. Acad. Sci. 98: 8668-8673. 11447280

Garnett, A. T., et al. (2009). Identification of direct T-box target genes in the developing zebrafish mesoderm. Development 136(5): 749-60. PubMed Citation: 19158186

Gofflot, F., Hall, M. and Morriss-Kay, G. M. (1997). Genetic patterning of the developing mouse tail at the time of posterior neuropore closure. Dev. Dyn. 210(4): 431-45. PubMed Citation: 9415428

Goldstein, R. E., et al. (1999). Huckebein repressor activity in Drosophila terminal patterning is mediated by Groucho. Development 126: 3747-3755. PubMed Citation: 10433905

Goto, H., Kimmey, S. C., Row, R. H., Matus, D. Q. and Martin, B. L. (2017). FGF and canonical Wnt signaling cooperate to induce paraxial mesoderm from tailbud neuromesodermal progenitors through regulation of a two-step EMT. Development [Epub ahead of print]. PubMed ID: 28242612

Glasgow, E., Karavanov, A. A. and Dawid, I. B. (1997). Neuronal and neuroendocrine expression of lim3, a LIM class homeobox gene, is altered in mutant zebrafish with axial signaling defects. Dev. Biol. 192(2): 405-419 . PubMed Citation: 9441677

Greenwood, S. and Struhl, G. (1997). Different levels of Ras activity can specify distinct transcriptional and morphological consequences in early Drosophila embryos. Development 124(23): 4879-4886. PubMed Citation: 9428424

Griffin, K., Patient, R. and Holder, N. (1995). Analysis of FGF function in normal and no tail zebrafish embryos reveals separate mechanisms for formation of the trunk and the tail. Development 121: 2983-2994. PubMed Citation: 7555724

Gross, J. M. and McClay, D. R. (2001). The role of Brachyury (T) during gastrulation movements in the sea urchin Lytechinus variegatus. Dev. Bio. 239: 132-147. PubMed Citation: 11784024

Harada, Y., Yasuo, H. and Satoh, N. (1995). A sea urchin homologue of the chordate Brachyury (T) gene is expressed in the secondary mesenchyme founder cells. Development 121: 2747-2754 . PubMed Citation: 7555703

Harrison, S. M. (2000). Sp5, a new member of the Sp1 family, is dynamically expressed during development and genetically interacts with Brachyury. Dev. Bio. 227: 358-372. PubMed Citation: 11071760

Harvey, S. A., et al. (2010). no tail integrates two modes of mesoderm induction. Development 137(7): 1127-35. PubMed Citation: 20215349

Hashimoto, H., Enomoto, A., Kumano, G. and Nishida, H. (2011). The transcription factor FoxB mediates temporal loss of cellular competence for notochord induction in ascidian embryos. Development 138(12): 2591-600. PubMed Citation: 21610035

Hotta, K., et al. (2000). Characterization of Brachyury-downstream notochord genes in the Ciona intestinalis embryo. PubMed Citation: 10898962 Dev. Bio. 224: 69-80.

Ikeda, T. and Satou, Y. (2016). Differential temporal control of Foxa.a and Zic-r.b specifies brain versus notochord fate in the ascidian embryo. Development [Epub ahead of print]. PubMed ID: 27888196

Isaacs, H. V., Pownall, M. E. and Slack, J. M. (1994). eFGF regulates Xbra expression during Xenopus gastrulation. EMBO J. 13(19): 4469-4481. PubMed Citation: 7925289

Jacob, A., Budhiraja, S., and Reichel, R. R. (1997). Differential induction of HNF-3 transcription factors during neuronal differentiation. Exp. Cell Res. 234(2): 277-284. PubMed Citation: 9260895

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. PubMed Citation: 10349616

Keller, R., Shih, J. and Domingo, C. (1992). The patterning and function of protrusive activity during convergence and extension of the Xenopus organizer. Development 1992 supplement: 81-91. PubMed Citation: 1299372

King. T., Beddington, R. S. P. and Brown, N. A. (1998). The role of the brachyury gene in heart development and left--right specification in the mouse. Mech. Dev. 79(1-2): 29-37. PubMed Citation: 10349618

Kispert, A., Herrmann, B.G., Leptin, M. and Reuter, R. (1994). Homologs of the mouse Brachyury gene are involved in the specification of posterior terminal structures in Drosophila, Tribolium, and Locusta. Genes Dev 8:2137-2150. PubMed Citation: 7958884

Kispert, A., Koschorz, B. and Herrmann, B. G. (1995a). The T protein encoded by Brachyury is a tissue-specific transcriptionn factor. EMBO J 14: 4763-4772. PubMed Citation: 7588606

Kispert, A., et al. (1995b). The chick Brachyury gene: developmental expression pattern and response to axial induction by localized activin. Dev Biol 168: 406-415. PubMed Citation: 7729577

Knezevic, V., De Santo, R. and Mackem, S. (1998). Continuing organizer function during chick tail development. Development 125(10): 1791-1801. PubMed Citation: 9550712

Kubo, A., et al. (2010). Genomic cis-regulatory networks in the early Ciona intestinalis embryo. Development 137(10): 1613-23. PubMed Citation: 20392745

Kumano, G. and Smith, W. C. (2000). FGF signaling restricts the primary blood islands to ventral mesoderm. Dev. Bio. 228: 304-314. PubMed Citation: 11112331

Kusch, T. and Reuter, R. (1999). Functions for Drosophila brachyenteron and forkhead in mesoderm specification and cell signalling. Development 126: 3991-4003. PubMed Citation: 10457009

Kusch, T., Storck, T., Walldorf, U. and Reuter, R. (2002). Brachyury proteins regulate target genes through modular binding sites in a cooperative fashion. Genes Dev. 16: 518-529. 11850413

Lartillot, N., et al. (2002). Expression pattern of Brachyury in the mollusc Patella vulgata suggests a conserved role in the establishment of the AP axis in Bilateria. Development 129: 1411-1421. 11880350

Latimer, A. J., et al. (2002). Delta-Notch signaling induces hypochord development in zebrafish. Development 129: 2555-2563. 12015285

Latinkic. B. V., et al. (1997). The Xenopus Brachyury promoter is activated by FGF and low concentrations of activin and suppressed by high concentrations of activin and by paired-type homeodomain proteins. Genes Dev. 11(23): 3265-3276. PubMed Citation: 9389657

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. PubMed Citation: 10079237

Lemaire, P., et al. (1998). A role for the vegetally expressed Xenopus gene Mix.1 in endoderm formation and in the restriction of mesoderm to the marginal zone. Development 125(13): 2371-2380. PubMed Citation: 9609820

Lerchner, W., et al. (2000). Region-specific activation of the Xenopus Brachyury promoter involves active repression in ectoderm and endoderm: a study using transgenic frog embryos. Development 127: 2729-2739. PubMed Citation: 10821770

Liu, C., et al. (2003). A role for the mesenchymal T-box gene Brachyury in AER formation during limb development. Development 130: 1327-1337. 12588849

Liu, X. and Lengyel. J. A. (2000). Drosophila arc encodes a novel adherens junction-associated PDZ domain protein required for wing and eye development. Dev. Biol. 221: 419-434. PubMed Citation: 10790336

Makita, R., et al. (1998). Zebrafish wnt11: pattern and regulation of the expression by the yolk cell and no tail activity. Mech. Dev. 71(1-2): 165-176. PubMed Citation: 10790336

Marcellini, S., et al. (2003). Evolution of Brachyury proteins: identification of a novel regulatory domain conserved within Bilateria. Dev. Biol. 260: 352-361. 12921737

Martin, B. L. and Kimelman, D. (2008). Regulation of canonical Wnt signaling by Brachyury is essential for posterior mesoderm formation. Dev. Cell 15(1): 121-33. PubMed Citation: 18606146

Martin, B. L. and Kimelman, D. (2010). Brachyury establishes the embryonic mesodermal progenitor niche. Genes Dev. 24(24): 2778-83. PubMed Citation: 21159819

Melby, A. E., Warga, R. M. and Kimmel, C. B. (1996). Specification of cell fates at the dorsal margin of the zebrafish gastrula. Development 122: 2225-2237. PubMed Citation: 8681803

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

Muller, C. W. and Herrmann, B. G. (1997). Crystallographic structure of the T domain-DNA complex of the Brachyury transcription factor. Nature 389: 884-888. PubMed Citation: 9349824

Murakami, R., Shigenaga, A., Kawakita, M., Takimoto,K., Yamaoka, I., Akasaka, K. and Shimada, H. (1995). aproctous, a locus that is necessary for the development of the proctodeum in Drosophila embryos, encodes a homolog of the vertebrate Brachyury gene. Roux's Arch Dev Biol 205:89-96

Nakatani, Y., et al. (1996). Basic fibroblast growth factor induces notochord formation and the expression of As-T, a Brachyury homolog, during ascidian embryogenesis. Development 122: 2023-2031. PubMed Citation: 8681783

Nishino, A., et al. (2001). Brachyury (T) gene expression and notochord development in Oikopleura longicauda (Appendicularia, Urochordata). Dev. Genes Evol. 211(5): 219-31. 11455437

Northrop, J. L.,and Kimelman, D. (1994). Dorsal-ventral differences in Xcad-3 expression in response to FGF-mediated induction in Xenopus. Dev Biol 161: 490-503. PubMed Citation: 7906234

Northrop, J., et al. (1995). BMP-4 regulates the dorsal-ventral differences in FGF/MAPKK-mediated mesoderm induction in Xenopus. Dev. Biol. 172(1): 242-252. PubMed Citation:

Okumura, T., Matsumoto, A., Tanimura, T. and Murakami, R. (2005). An endoderm-specific GATA factor gene, dGATAe, is required for the terminal differentiation of the Drosophilaendoderm. Dev. Biol. 278(2): 576-86. 15680371

O'Reilly, M. A., Smith, J. C. and Cunliffe, V. (1995). Patterning of the mesoderm in Xenopus: dose-dependent and synergistic effects of Brachyury and Pintallavis. Development 121: 1351-1359. PubMed Citation: 7789266

Panitz, F., et al. (1998). The Spemann organizer-expressed zinc finger gene Xegr-1 responds to the MAP kinase/Ets-SRF signal transduction pathway. EMBO J. 17: 4414-4425. PubMed Citation: 9687509

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

Papin, C., van Grunsven, L. A., Verschueren, K., Huylebroeck, D. and Smith, J. C. (2002). Dynamic regulation of Brachyury expression in the amphibian embryo by XSIP1. Mech. Dev. 111(1-2): 37-46. 11804777

Passamaneck, Y. J., et al. (2009). Direct activation of a notochord cis-regulatory module by Brachyury and FoxA in the ascidian Ciona intestinalis. Development 136(21): 3679-89. PubMed Citation: 19820186

Peterson, K. J., et al. (1999a). A comparative molecular approach to mesodermal patterning in basal deuterostomes: the expression pattern of Brachyury in the enteropneust hemichordate Ptychodera flava. Development 126(1): 85-95. PubMed Citation: 9834188

Peterson, K. J., et al. (1999b), Expression pattern of Brachyury and Not in the sea urchin: comparative implications for the origins of mesoderm in the basal deuterostomes. Dev. Biol. 207(2): 419-31. PubMed Citation: 10068473

Poeck, B., Balles, J. and Pflugfelder, G.O. (1993). Transcript identification in the optomotor-blind locus of Drosophila melanogaster by intragenic recombination mapping and PCR-aided sequence analysis of lethal point mutations. Mol. Gen. Genet. 238(3): 325-32. PubMed Citation: 8492800

Rashbass, P., et al. (1994). Alterations in gene expression during mesoderm formation and axial patterning in Brachyury (T) embryos. Int. J. Dev. Biol. 38: 35-44. PubMed Citation: 7915533

Rast, J. P., et al. (2002). brachyury target genes in the early sea urchin embryo isolated by differential macroarray screening. Dev. Biol. 246: 191-208. 12027442

Rennebeck, G., et al. (1998). Mouse brachyury the second (T2) is a gene next to classical T and a candidate gene for tct. Genetics 150(3): 1125-31. PubMed Citation: 9799264

Saka, Y., Tada, M. and Smith, J. C. (2000). A screen for targets of the Xenopus T-box gene Xbra. Mech. Dev. 93: 27-39. PubMed Citation: 10781937

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: 843-853. PubMed Citation: 7600961

Schulte-Merker, S. and Smith, J. C. (1995). Mesoderm formation in response to Brachyury requires FGF signalling. Curr. Biol. 5(1):62-67. PubMed Citation: 7535172

Shinga, J., et al. (2001). Early patterning of the prospective midbrain-hindbrain boundary by the HES-related gene XHR1 in Xenopus embryos. Mech. Dev. 109(2): 225-39. 11731236

Shoguchi, E., Satoh, N. and Maruyama, Y. K. (1999). Pattern of Brachyury gene expression in starfish embryos resembles that of hemichordate embryos but not of sea urchin embryos. Mech. Dev. 82(1-2): 185-189. PubMed Citation: 10354483

Simeone, A., D'Apice, M.R., Nigro, V., Casanova, J., Graziani, F., Acampora, D. and Avantaggiato, V. (1994). Orthopedia, a novel homeobox-containing gene expressed in the developing CNS of both mouse and Drosophila. Neuron 13: 83-101. 7913821

Singer, J. B., et al. (1996). Drosophila brachyenteron regulates gene activity and morphogenesis in the gut. Development 122: 3703-18 . PubMed Citation: 9012492

Spring, J., et al. (2002). Conservation of Brachyury, Mef2, and Snail in the myogenic lineage of jellyfish: A connection to the mesoderm of bilateria. Dev. Biol. 244: 372-384. 11944944

Strong, C. F., et al. (2000). Xbra3 induces mesoderm and neural tissue in Xenopus laevis. Dev. Biol. 222: 405-419. PubMed Citation: 10837128

Tada, M. and Smith, J. C. (2000). Xwnt11 is a target of Xenopus Brachyury: regulation of gastrulation movements via Dishevelled, but not through the canonical Wnt pathway. Development 127: 2227-2238 . PubMed Citation: 10769246

Taira, M., Saint-Jeannet, J.-P. and Dawid, I. B. (1997). Role of the Xlim-1 and Xbra genes in anteroposterior patterning of neural tissue by the head and trunk organizer. Proc. Natl. Acad. Sci. 94: 895-900. PubMed Citation: 9023353

Takahashi, H., et al. (1999). Evolutionary alterations of the minimal promoter for notochord-specific Brachyury expression in ascidian embryos. Development 126: 3725-3734. PubMed Citation: 10433903

Teillet, M. A., Lapointe, F., Le Douarin, N. M. (1998). The relationships between notochord and floor plate in vertebrate development revisited. Proc. Natl. Acad. Sci. 95(20): 11733-8. PubMed Citation: 10433903

Terazawa, K. and Satoh. N. (1997). Formation of the chordamesoderm in the amphioxus embryo: Analysis with Brachyury and fork head/HNF-3 genes. Dev. Genes Evol. 207: 1-11 . PubMed Citation: 20607475

Umbhauer, M., et al. (1994). Control of somitic expression of tenascin in Xenopus embryos by myogenic factors and Brachyury. Dev. Dyn. 200 (4): 269-277. PubMed Citation: 7527682

Viebahn, C., et al. (2002). Low proliferative and high migratory activity in the area of Brachyury expressing mesoderm progenitor cells in the gastrulating rabbit embryo. Development 129: 2355-2365. 11973268

Vonica, A., and Gumbiner, B. M. (2002). Zygotic wnt activity is required for Brachyury expression in the early Xenopus laevis embryo. Dev. Bio. 250: 112-127. 12297100

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

Wilson, V., et al. (1995). The T gene is necessary for normal mesodermal morphogenetic cell movements during gastrulation. Development 121: 877-886. PubMed Citation: 7720590"> 7720590

Wilson, V. and Beddington, R. (1997). Expression of T protein in the primitive streak is necessary and sufficient for posterior mesoderm movement and somite differentiation. Dev. Biol. 192(1): 45-58. PubMed Citation: 9405096

Woollard, A. and Hodgkin, J. (2000). The Caenorhabditis elegans fate-determining gene mab-9 encodes a T-box protein required to pattern the posterior hindgut. Genes Dev. 14: 596-603. PubMed Citation: 10716947

Wu, L. H. and Lengyel, J. A. (1998). Role of caudal in hindgut specification and gastrulation suggests homology between Drosophila amnioproctodeal invagination and vertebrate blastopore. Development 125: 2433-2442. PubMed Citation: 9609826

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

Yamada, T. (1994). Caudalization by the amphibian organizer: brachyury, convergent extension and retinoic acid. Development 120: 3051-62. PubMed Citation:

Yasuo, H. and Satoh, N. (1998). Conservation of the developmental role of Brachyury in notochord formation in a Urochordate, the ascidian Halocynthia roretzi. Dev. Biol. 200(2): 158-170 . PubMed Citation: 9705224

Yasuoka, Y., Shinzato, C. and Satoh, N. (2016). The mesoderm-forming gene brachyury regulates ectoderm-endoderm demarcation in the coral Acropora digitifera. Curr Biol [Epub ahead of print]. PubMed ID: 27693135

Zhang, S., Holland, N. D. and Holland, L. Z. (1997). Topographic changes in nascent and early mesoderm in amphioxus embryos studied by DiI labeling and by in situ hybridization for a Brachyury gene. Dev. Genes Evol. 206: 532-535 . PubMed Citation:

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. PubMed Citation: 9441683

Brachyenteron/T-related gene: Biological Overview | Evolutionary Homologs | Regulation | Developmental Biology | Effects of Mutation

date revised: 12 December 2016

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

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