Gene Symbol: fog
Description: folded gastrulation
Alias: 114, CG9559, CR41108, CR41109, CR41110, Dmel\CG9559, Fog, M67, l(1)114, l(1)16-3-190, l(1)20Ba, l114, folded gastrulation, CG9559-PA, CG9559-PB, CG9559-PC, CG9559-PD, CG9559-PE, CG9559-PF, fog-PA, fog-PB, fog-PC, fog-PD, fog-PE, fog-PF
Species: fruit fly
Products:     fog

Top Publications

  1. Bertet C, Sulak L, Lecuit T. Myosin-dependent junction remodelling controls planar cell intercalation and axis elongation. Nature. 2004;429:667-71 pubmed
    ..This simple cellular mechanism provides a general model for polarized morphogenesis in epithelial organs. ..
  2. Costa M, Wilson E, Wieschaus E. A putative cell signal encoded by the folded gastrulation gene coordinates cell shape changes during Drosophila gastrulation. Cell. 1994;76:1075-89 pubmed
    The folded gastrulation (fog) gene is required during Drosophila gastrulation for two morphogenetic movements, formation of the ventral furrow and invagination of the posterior midgut primordium...
  3. Sweeton D, Parks S, Costa M, Wieschaus E. Gastrulation in Drosophila: the formation of the ventral furrow and posterior midgut invaginations. Development. 1991;112:775-89 pubmed
    ..Mutations in the concertina or folded gastrulation genes appear to block the transition to the second phase in both the ventral furrow and the posterior ..
  4. Dawes Hoang R, Parmar K, Christiansen A, Phelps C, Brand A, Wieschaus E. folded gastrulation, cell shape change and the control of myosin localization. Development. 2005;132:4165-78 pubmed
    ..In this paper, we analyze the function of the folded gastrulation gene (fog), a known target of the patterning gene twist...
  5. Barrett K, Leptin M, Settleman J. The Rho GTPase and a putative RhoGEF mediate a signaling pathway for the cell shape changes in Drosophila gastrulation. Cell. 1997;91:905-15 pubmed also presented that DRhoGEF2 mediates these specific cell shape changes in response to the extracellular ligand, Fog. Together, these results establish a Rho-mediated signaling pathway that is essential for the major morphogenetic ..
  6. Morize P, Christiansen A, Costa M, Parks S, Wieschaus E. Hyperactivation of the folded gastrulation pathway induces specific cell shape changes. Development. 1998;125:589-97 pubmed
    ..In embryos mutant for folded gastrulation (fog), these cell shape changes occur but the timing and synchrony of the constrictions are abnormal...
  7. Pouille P, Ahmadi P, Brunet A, Farge E. Mechanical signals trigger Myosin II redistribution and mesoderm invagination in Drosophila embryos. Sci Signal. 2009;2:ra16 pubmed publisher
    ..The second wave is controlled by the transcription factor Twist and requires the secreted protein Fog. Complete mesoderm invagination requires redistribution of the motor protein Myosin II to the apical side of the ..
  8. Wieschaus E. Embryonic transcription and the control of developmental pathways. Genetics. 1996;142:5-10 pubmed
  9. Nikolaidou K, Barrett K. A Rho GTPase signaling pathway is used reiteratively in epithelial folding and potentially selects the outcome of Rho activation. Curr Biol. 2004;14:1822-6 pubmed member that is thought to activate DRhoGEF2 and is itself activated by a putative ligand, Folded gastrulation (Fog)...

More Information


  1. Leptin M. Drosophila gastrulation: from pattern formation to morphogenesis. Annu Rev Cell Dev Biol. 1995;11:189-212 pubmed
    ..These genes code for transcription factors that regulate cell behavior by controlling expression of target genes whose products modulate the cytoskeleton and other subcellular compartments. ..
  2. Miklos G, Cotsell J. Chromosome structure at interfaces between major chromatin types: alpha- and beta-heterochromatin. Bioessays. 1990;12:1-6 pubmed
    ..The recent microcloning of part of this region, the isolation of a monoclonal antibody to a beta-heterochromatin binding protein, and new in situ studies now shed a little more light on this chromosomal region. ..
  3. Martin A, Gelbart M, Fernandez Gonzalez R, Kaschube M, Wieschaus E. Integration of contractile forces during tissue invagination. J Cell Biol. 2010;188:735-49 pubmed publisher
    ..Thus, pulsed actomyosin contractions require a supracellular, tensile meshwork to transmit cellular forces to the tissue level during morphogenesis. ..
  4. Johnson T, Moore K, Whisstock J, Warr C. Maternal Torso-Like Coordinates Tissue Folding During Drosophila Gastrulation. Genetics. 2017;206:1459-1468 pubmed publisher
    ..In addition, loss of tsl strongly suppressed the effects of ectopic expression of Folded Gastrulation (Fog), a secreted protein that promotes apical constriction...
  5. Perrimon N, Smouse D, Miklos G. Developmental genetics of loci at the base of the X chromosome of Drosophila melanogaster. Genetics. 1989;121:313-31 pubmed
    ..There are 22 loci which, when mutated, are zygotic lethals and three of these, legless/runt, folded gastrulation and 13E3, have severe effects on embryonic development...
  6. Wu L, Lengyel J. Role of caudal in hindgut specification and gastrulation suggests homology between Drosophila amnioproctodeal invagination and vertebrate blastopore. Development. 1998;125:2433-42 pubmed
    ..These effects are achieved by the function of caudal in activating different target genes, namely folded gastrulation, which is required for invagination of the posterior gut primordium, and fork head and wingless, which are ..
  7. Lammel U, Saumweber H. X-linked loci of Drosophila melanogaster causing defects in the morphology of the embryonic salivary glands. Dev Genes Evol. 2000;210:525-35 pubmed
    ..From these, 39 lines could be assigned to nine complementation groups: armadillo, brinker, folded gastrulation, giant, hindsight, Notch, runt, stardust and twisted gastrulation.
  8. Ratnaparkhi A. Signaling by Folded gastrulation is modulated by mitochondrial fusion and fission. J Cell Sci. 2013;126:5369-76 pubmed publisher
    Mitochondria are increasingly being identified as integrators and regulators of cell signaling pathways. Folded gastrulation (Fog) is a secreted signaling molecule best known for its role in regulating cell shape change at the ventral ..
  9. Lifschytz E. Fine-Structure Analysis and Genetic Organization at the Base of the X Chromosome in DROSOPHILA MELANOGASTER. Genetics. 1978;88:457-67 pubmed
    ..Section 20 is considered to be a transition region between the mostly heterochromatic and mostly euchromatic regions of the X chromosome; the differences between them are suggested as being merely quantitative. ..
  10. Johnson T, Henstridge M, Herr A, Moore K, Whisstock J, Warr C. Torso-like mediates extracellular accumulation of Furin-cleaved Trunk to pattern the Drosophila embryo termini. Nat Commun. 2015;6:8759 pubmed publisher
    ..Our data represent an alternative mechanism for the spatial control of receptor signalling, and define a different role for perforin-like proteins in eukaryotes. ..
  11. Yip M, Lamka M, Lipshitz H. Control of germ-band retraction in Drosophila by the zinc-finger protein HINDSIGHT. Development. 1997;124:2129-41 pubmed
    ..Second, hindsight might function both to maintain the amnioserosa and to regulate chemical signaling from the amnioserosa to the epidermal ectoderm, thus coordinating the cell shape changes and movements that drive germ-band retraction...
  12. Driquez B, Bouclet A, Farge E. Mechanotransduction in mechanically coupled pulsating cells: transition to collective constriction and mesoderm invagination simulation. Phys Biol. 2011;8:066007 pubmed publisher
    ..invagination at the onset of gastrulation was proposed to be triggered in response to the activation of the Fog mechanotransduction pathway by the Snail-dependent active mechanical oscillations of cell apex sizes...
  13. Pouille P, Farge E. Hydrodynamic simulation of multicellular embryo invagination. Phys Biol. 2008;5:015005 pubmed publisher
  14. Mathew S, Rembold M, Leptin M. Role for Traf4 in polarizing adherens junctions as a prerequisite for efficient cell shape changes. Mol Cell Biol. 2011;31:4978-93 pubmed publisher
    ..Traf4 acts in a TNF receptor- and Jun N-terminal protein kinase (JNK)-independent manner to fine-tune the assembly of adherens junctions in the invaginating mesodermal cells...
  15. Ratnaparkhi A, Zinn K. The secreted cell signal Folded Gastrulation regulates glial morphogenesis and axon guidance in Drosophila. Dev Biol. 2007;308:158-68 pubmed
    ..This process is affected in mutant embryos lacking zygotic function of the folded gastrulation (fog) gene, which encodes a putative secreted protein...
  16. Chovnick A, Finnertty V, Schalet A, Duck P. Studies on genetic organization in higher organisms. I. Analysis of a complex gene in Drosophila melanogaster. Genetics. 1969;62:145-60 pubmed
  17. Leptin M. Gastrulation in Drosophila: the logic and the cellular mechanisms. EMBO J. 1999;18:3187-92 pubmed
  18. Bard J. Epithelial rearrangement and Drosophila gastrulation. Bioessays. 1991;13:409-11 pubmed
  19. de Renzis S, Elemento O, Tavazoie S, Wieschaus E. Unmasking activation of the zygotic genome using chromosomal deletions in the Drosophila embryo. PLoS Biol. 2007;5:e117 pubmed
    ..We propose that this regulatory mode links morphogen gradients with temporal regulation during the maternal-to-zygotic transition. ..
  20. Manning A, Peters K, Peifer M, Rogers S. Regulation of epithelial morphogenesis by the G protein-coupled receptor mist and its ligand fog. Sci Signal. 2013;6:ra98 pubmed publisher
    ..nucleotide-binding protein)-coupled receptor (GPCR) that transduces signals from the secreted ligand Folded gastrulation (Fog) in cultured cells...
  21. Paradi E, Vogel E, Szilagyi E. Effect of storage and dose on MMS-induced deletions. Complementation analysis of X-chromosomal recessive lethals in the zeste-white and maroon-like regions of Drosophila melanogaster. Mutat Res. 1983;111:145-59 pubmed
    ..An interesting aspect for further analysis is the apparent infrequency in the zeste-white region of alkylation-induced chromosomal breakage, as observed by various investigators for MMS, EMS and MNNG. ..
  22. Nie J, Mahato S, Zelhof A. The actomyosin machinery is required for Drosophila retinal lumen formation. PLoS Genet. 2014;10:e1004608 pubmed publisher
  23. Zusman S, Wieschaus E. Requirements for zygotic gene activity during gastrulation in Drosophila melanogaster. Dev Biol. 1985;111:359-71 pubmed
    Mutations at the folded gastrulation (fog) and twisted gastrulation (tsg) loci interfere with early morphogenetic movements in Drosophila melanogaster...
  24. Weng M, Wieschaus E. Myosin-dependent remodeling of adherens junctions protects junctions from Snail-dependent disassembly. J Cell Biol. 2016;212:219-29 pubmed publisher
    ..Our findings provide in vivo evidence for mechanosensitivity of cell-cell junctions and imply that myosin-mediated tension can prevent Snail-driven EMT. ..
  25. Butler L, Blanchard G, Kabla A, Lawrence N, Welchman D, Mahadevan L, et al. Cell shape changes indicate a role for extrinsic tensile forces in Drosophila germ-band extension. Nat Cell Biol. 2009;11:859-64 pubmed publisher
    ..We propose that cell shape change contributing to germ-band extension is a passive response to mechanical forces caused by the invaginating mesoderm. ..
  26. Oda H, Tsukita S, Takeichi M. Dynamic behavior of the cadherin-based cell-cell adhesion system during Drosophila gastrulation. Dev Biol. 1998;203:435-50 pubmed
    ..Moreover, the processes responsible for loss of epithelial cell polarity and elimination of preexisting DE-cadherin-based epithelial junctions during early mesodermal morphogenesis are discussed. ..
  27. Krasnow M. Genes that control organ form: lessons from bone and branching morphogenesis. Cold Spring Harb Symp Quant Biol. 1997;62:235-40 pubmed
  28. Wolfner M, Goldberg M. Harnessing the power of Drosophila genetics. Methods Cell Biol. 1994;44:33-80 pubmed
  29. Leptin M. Morphogenesis. Control of epithelial cell shape changes. Curr Biol. 1994;4:709-12 pubmed
    Cell-shape changes in the gastrulating Drosophila embryo generate two epithelial invaginations. These changes are regulated by four transcription factors, and folded gastrulation is a candidate downstream effector.
  30. Miklos G, Kelly L, Coombe P, Leeds C, Lefevre G. Localization of the genes shaking-B, small optic lobes, sluggish-A, stoned and stress-sensitive-C to a well-defined region on the X-chromosome of Drosophila melanogaster. J Neurogenet. 1987;4:1-19 pubmed
    ..belong to adjacent complementation groups, between lethals W2 and A112 near band 19F4; and stonedts1 and stress-sensitive-C1 are both alleles of the 8P1 lethal complementation group between lethals 114 and 13E3 near bands 20B-C.
  31. Peters K, Rogers S. Drosophila Ric-8 interacts with the G?12/13 subunit, Concertina, during activation of the Folded gastrulation pathway. Mol Biol Cell. 2013;24:3460-71 pubmed publisher
    ..We also use this system to conduct a structure-function analysis of Ric-8 and identify key residues required for both Cta interaction and cellular contractility. ..
  32. Fuse N, Yu F, Hirose S. Gprk2 adjusts Fog signaling to organize cell movements in Drosophila gastrulation. Development. 2013;140:4246-55 pubmed publisher
    ..melanogaster proceeds through sequential cell movements: ventral mesodermal (VM) cells are induced by secreted Fog protein to constrict their apical surfaces to form the ventral furrow, and subsequently lateral mesodermal (LM) ..
  33. Urbansky S, González Avalos P, Wosch M, Lemke S. Folded gastrulation and T48 drive the evolution of coordinated mesoderm internalization in flies. elife. 2016;5: pubmed publisher
    ..Here we identify two genes, folded gastrulation and t48, which in the evolution of fly gastrulation acted as a likely switch from an ingression ..
  34. Rahman R, Lindsley D. Male-sterilizing interactions between duplications and deficiencies for proximal X-chromosome material in Drosophila melanogaster. Genetics. 1981;99:49-64 pubmed
    ..We conclude that there is no single factor at the base of the X that is required for the fertility of males whose genotype is otherwise normal. ..
  35. Anaka M, Lynn A, McGinn P, Lloyd V. Genomic imprinting in Drosophila has properties of both mammalian and insect imprinting. Dev Genes Evol. 2009;219:59-66 pubmed publisher
  36. Wieschaus E, Nusslein Volhard C, Kluding H. Krüppel, a gene whose activity is required early in the zygotic genome for normal embryonic segmentation. Dev Biol. 1984;104:172-86 pubmed
  37. Bergman C, Quesneville H, Anxolabehere D, Ashburner M. Recurrent insertion and duplication generate networks of transposable element sequences in the Drosophila melanogaster genome. Genome Biol. 2006;7:R112 pubmed
    ..Complex relationships of TE nesting in beta-heterochromatic regions raise the possibility of a co-suppression network that may act as a global surveillance system against the majority of TE families in D. melanogaster. ..
  38. Chung S, Kim S, Andrew D. Uncoupling apical constriction from tissue invagination. elife. 2017;6: pubmed publisher
    ..Using Drosophila salivary gland (SG) invagination as a model, we show that regulation of folded gastrulation expression by the Fork head transcription factor is required for apicomedial accumulation of Rho kinase ..
  39. Muller H, Wieschaus E. armadillo, bazooka, and stardust are critical for early stages in formation of the zonula adherens and maintenance of the polarized blastoderm epithelium in Drosophila. J Cell Biol. 1996;134:149-63 pubmed
    ..These results suggest that early stages in the assembly of the ZA are critical for the stability of the polarized blastoderm epithelium...
  40. Kerridge S, Munjal A, Philippe J, Jha A, de Las Bayonas A, Saurin A, et al. Modular activation of Rho1 by GPCR signalling imparts polarized myosin II activation during morphogenesis. Nat Cell Biol. 2016;18:261-70 pubmed publisher
    ..We propose that GPCR and G proteins constitute a general pathway for controlling actomyosin contractility in epithelia and that the activity of this pathway is polarized by tissue-specific regulators. ..
  41. Huang A, Rusch J, Levine M. An anteroposterior Dorsal gradient in the Drosophila embryo. Genes Dev. 1997;11:1963-73 pubmed
    ..This study also provides evidence that neurogenic repressors may be important for the establishment of the sharp mesoderm/neuroectoderm boundary in the early embryo. ..
  42. Seher T, Narasimha M, Vogelsang E, Leptin M. Analysis and reconstitution of the genetic cascade controlling early mesoderm morphogenesis in the Drosophila embryo. Mech Dev. 2007;124:167-79 pubmed
    ..Twist, which acts at the top of the mesodermal developmental cascade in Drosophila, two targets, Snail and Fog, are known to be necessary for the first morphogenetic event, the orderly invagination of the mesoderm...
  43. Andrews J, Smith M, Merakovsky J, Coulson M, Hannan F, Kelly L. The stoned locus of Drosophila melanogaster produces a dicistronic transcript and encodes two distinct polypeptides. Genetics. 1996;143:1699-711 pubmed
    ..The protein encoded by the first ORF contains four tandemly repeated motifs, and one domain of the protein encoded by the second ORF shows similarity to a family of proteins (AP50s) associated with clathrin assembly protein complexes. ..
  44. Knust E, Muller H. Drosophila morphogenesis: orchestrating cell rearrangements. Curr Biol. 1998;8:R853-5 pubmed
    ..Recent results have shown that, during Drosophila gastrulation, multiple signalling pathways act to orchestrate the complex cell rearrangements. ..
  45. Fernandez Sanchez M, Serman F, Ahmadi P, Farge E. Mechanical induction in embryonic development and tumor growth integrative cues through molecular to multicellular interplay and evolutionary perspectives. Methods Cell Biol. 2010;98:295-321 pubmed publisher
    ..gastrulation, through mechanotransduction processes involving the Armadillo/beta-catenin and the downstream of Fog Rho pathways...
  46. Strecker T, Yip M, Lipshitz H. Genetic control of cell fate in the termini of the Drosophila embryo. Dev Biol. 1992;150:422-6 pubmed
    ..Genes that control dorsoventral pattern in the termini and genes that program terminal morphogenesis act in distinct branches of the genetic hierarchy that are independent of tailless. ..
  47. Lammel U, Meadows L, Saumweber H. Analysis of Drosophila salivary gland, epidermis and CNS development suggests an additional function of brinker in anterior-posterior cell fate specification. Mech Dev. 2000;92:179-91 pubmed
    ..Furthermore, we observed a preferential loss of lateral neuroblasts in the anterior parasegment. Together, these phenotypes suggest that brinker not only plays a role in dorso-ventral but also in anterior-posterior axis patterning. ..
  48. Lifschytz E, Yakobovitz N. The role of X-linked lethal and viable male-sterile mutations in male gametogenesis of Drosophila melanogaster: genetic analysis. Mol Gen Genet. 1978;161:275-84 pubmed
    ..It is further shown that about the same proportion of vital genes is operating in female gonads and that the two groups overlap by about 70%. The role of viable and lethal male gametogenesis is discussed in detail. ..
  49. Tepass U, Hartenstein V. Epithelium formation in the Drosophila midgut depends on the interaction of endoderm and mesoderm. Development. 1994;120:579-90 pubmed
  50. Oda H, Tsukita S. Real-time imaging of cell-cell adherens junctions reveals that Drosophila mesoderm invagination begins with two phases of apical constriction of cells. J Cell Sci. 2001;114:493-501 pubmed
    ..Furthermore, we showed that the mesoderm invagination of folded gastrulation mutant embryos lacked the normal two constriction phases, and instead began with asynchronous, feeble cell ..
  51. Fabrowski P, Necakov A, Mumbauer S, Loeser E, Reversi A, Streichan S, et al. Tubular endocytosis drives remodelling of the apical surface during epithelial morphogenesis in Drosophila. Nat Commun. 2013;4:2244 pubmed publisher
    ..These data collectively demonstrate a requirement for endocytosis in morphogenetic remodelling during epithelial development. ..
  52. Miklos G, de Couet H. The mutations previously designated as flightless-I3, flightless-O2 and standby are members of the W-2 lethal complementation group at the base of the X-chromosome of Drosophila melanogaster. J Neurogenet. 1990;6:133-51 pubmed
    ..These viable flightless mutations are all hypomorphs since the homozygous deficiency of the W-2 X-chromosomal region is lethal to the organism. ..
  53. Leptin M, Casal J, Grunewald B, Reuter R. Mechanisms of early Drosophila mesoderm formation. Dev Suppl. 1992;:23-31 pubmed
    ..The events that lead to the cell shape changes mediating ventral furrow formation require the transcription of zygotic genes under the control of twist and snail. Such genes can be isolated by molecular and genetic screens. ..
  54. Okegbe T, Dinardo S. The endoderm specifies the mesodermal niche for the germline in Drosophila via Delta-Notch signaling. Development. 2011;138:1259-67 pubmed publisher
    ..Because in mammals primordial germ cells also track through endoderm on their way to the genital ridge, our work raises the possibility that conserved mechanisms are employed to regulate germline niche formation...
  55. Kolesnikov T, Beckendorf S. 18 wheeler regulates apical constriction of salivary gland cells via the Rho-GTPase-signaling pathway. Dev Biol. 2007;307:53-61 pubmed
    ..interactions and double mutant analysis suggest that 18W affects the Rho-GTPase-signaling pathway not through Fog and RhoGEF2, but rather by inhibiting Rho GTPase activating proteins (RhoGAPs)...
  56. Hemavathy K, Hu X, Ashraf S, Small S, Ip Y. The repressor function of snail is required for Drosophila gastrulation and is not replaceable by Escargot or Worniu. Dev Biol. 2004;269:411-20 pubmed
    ..Together, the results show that the repressor function of Snail is essential for gastrulation. The repression of target genes by Snail may permit other factors in the ventral cells to positively promote mesoderm invagination. ..
  57. Fuse N, Hisata K, Katzen A, Matsuzaki F. Heterotrimeric G proteins regulate daughter cell size asymmetry in Drosophila neuroblast divisions. Curr Biol. 2003;13:947-54 pubmed
    ..Furthermore, the multiple equal cleavages of G beta mutant neuroblasts accompany neural defects; this finding suggests indispensable roles of eccentric division in assuring the stem cell properties of neuroblasts. ..
  58. Myat M, Andrew D. Organ shape in the Drosophila salivary gland is controlled by regulated, sequential internalization of the primordia. Development. 2000;127:679-91 pubmed
    ..We propose that fas is required for hkb-dependent signaling events that coordinate internalization. ..