
en expression is diminished in wg and arm mutants. An allelic series was described for each with with increasingly severe segment polarity phenotypes and earlier and more severe loss of en expression. The segmental stripes decay with a similar asymmetric pattern in wg and arm mutants. (Peifer M., 1991)
en expression was observed sequentially in 5 'centers' anterior to the mandibular segment starting at embryonic stage 8.(Schmidt-Ott U., 1992)
The en protein is expressed in a specified subset of neuroblasts in embryonic stages 8-11. (Doe C.Q., 1992)
Delayed changes in en expression were observed in response to ectopic
eve expression in eve[hs.PS] embryos. Four different expression patterns
were observed depending on the timing of ectopic eve protein induction.
(Manoukian A.S., 1992)
| Regulatory element | Localisation of element | Trans-regulatory factor | References |
| engrailed 5'-flank | promoter region | HUNCHBACK | Zuo et al., 1991 |

Engrailed-mediated repression of Ultrabithorax is necessary for the parasegment 6 identity in Drosophila. Development 120: 3205-3212 (1994)
Genetic factors controlling the expression of the abdominal-A gene of Drosophila within its domain. Mech Dev 46: 15-25 (1994)
Regulation of polyhomeotic transcription may involve local changes in chromatin activity in Drosophila. Mech Dev 52: 343-355 (1995)
polyhomeotic appears to be a target of engrailed regulation in Drosophila. Development 121: 1691-1703 (1995)
Drawing a stripe in Drosophila imaginal disks: negative regulation of decapentaplegic and patched expression by engrailed. Genetics 139: 745-756 (1995)
Sequential organizing activities of engrailed, hedgehog and decapentaplegic in the Drosophila wing. Development 121: 2265-2278 (1995)
Selection and characterization of sequences with high affinity for the engrailed protein of Drosophila. Mech. Dev. 53: 185-195 (1995)
Selection and characterization of sequences with high affinity for the engrailed protein of Drosophila. Mech. Dev. 53: 185-195 (1995)
Autocatalysis and phenotypic expression of Drosophila homeotic gene Deformed: its dependence on polarity and homeotic gene function. Development 116: 1059-1068 (1992)
Drawing a stripe in Drosophila imaginal disks: negative regulation of decapentaplegic and patched expression by engrailed. Genetics 139: 745-756 (1995)
Segmentation and specification of the Drosophila mesoderm. Genes Dev 10: 3183-3194 (1996)
Effects of ectopic expression of caudal during Drosophila development. Development 109: 271-277 (1990)
A molecular gradient in early Drosophila embryos and its role in specifying the body pattern. Macdonald P; Struhl B. Nature 324: 537-545 (1986)
Early even-skipped stripes act as morphogenetic gradients at the single cell level to establish engrailed expression. Development 121: 4371-4382 (1995)
Concentration-dependent activities of the even-skipped protein in Drosophila embryos. Genes Dev 6: 1740-1751 (1992)
Control of segmental asymmetry in Drosophila embryos. Development 118: 785-796 (1993)
Components of wingless signalling in Drosophila. Nature 367: 76-80 (1994)
Stripe-specific regulation of pair-rule genes by hopscotch, a putative Jak family tyrosine kinase in Drosophila. Genes Dev 8: 300-312 (1994)
groucho and hedgehog regulate engrailed expression in the anterior compartment of the Drosophila wing. Development 121: 3467-3476 (1995)
Drosophila wingless sustains engrailed expression only in adjoining cells: evidence from mosaic embryos. Cell 77: 909-915 (1994)
Trithorax is required to maintain engrailed expression in a subset of engrailed-expressing cells. Mech Dev 52: 89-98 (1995)
The single-minded gene of Drosophila is required for the expression of genes important for the development of CNS midline cells. Cell 63: 63-75 (1990)
Activation and repression of transcription by the gap proteins hunchback and Kruppel in cultured Drosophila cells. Genes Dev 5: 254-264 (1991)
Activation and repression of transcription by the gap proteins hunchback and Kruppel in cultured Drosophila cells. Genes Dev 5: 254-264 (1991)
Evolution of distinct developmental functions of three Drosophila genes by acquisition of different cis-regulatory regions. Nature, 367: 83-87 (1994)
Complex regulation of early paired expression: initial activation by gap genes and pattern modulation by pair-rule genes. Development 117: 609-623 (1993)
Establishing parasegments in Drosophila embryos: roles of the odd-skipped and naked genes. Dev Biol 169: 295-308 (1995)
_
Establishing parasegments in Drosophila embryos: roles of the odd-skipped and naked genes. Dev Biol 169: 295-308 (1995)
Control of segmental asymmetry in Drosophila embryos. Development 118: 785-796 (1993)
Early even-skipped stripes act as morphogenetic gradients at the single cell level to establish engrailed expression. Development 121: 4371-4382 (1995)
Localized expression of sloppy paired protein maintains the polarity of Drosophila parasegments. Genes Dev 8: 899-913 (1994)
odd-paired: a zinc finger pair-rule protein required for the timely activation of engrailed and wingless in Drosophila embryos. Genes Dev 8: 105-117 (1994)
Evolution of distinct developmental functions of three Drosophila genes by acquisition of different cis-regulatory regions. Nature, 367: 83-87 (1994)
The function of engrailed and the specification of Drosophila wing pattern. Development 121: 3447-3456 (1995)
The Drosophila cubitus interruptus protein and its role in the wingless and hedgehog signal transduction pathways. Mech Dev 52: 137-150 (1995)
Control of segmental asymmetry in Drosophila embryos. Development 118: 785-796 (1993)
Pattern formation in Drosophila head development: the role of the orthodenticle homeobox gene. Development 121: 3561-3572 (1995)
tramtrack is a transcriptional repressor required for cell fate determination in the Drosophila eye. Genes Dev 7: 1085-1096(1993)
Comparative analysis of Engrailed-1 and Wnt-1 expression in the developing central nervous system of Xenopus laevis. Int J Dev Biol 38: 623-632 (1994)
Drawing a stripe in Drosophila imaginal disks: negative regulation of decapentaplegic and patched expression by engrailed. Genetics 139: 745-756 (1995)
Baumgartner, S., Martin, D., Hagios, C. and Chiquet-Ehrismann, R. (1994). ten[m], a Drosophila gene related to tenascin, is a new pair-rule gene. EMBO J. 13: 3728-3740
Binari, R. and Perrimon, N. (1994). Stripe-specific regulation of pair-rule genes by hopscotch, a putative Jak family tyrosine kinase in Drosophila. Genes Dev 8: 300-12
Bourbon,H. M., Martin-Blanco E., Rosen D. and Kornberg ,T. B. (1995). Phosphorylation of the Drosophila engrailed protein at a site outside its homeodomain enhances DNA binding. J Biol Chem 270: 11130-11139
Breen, T. R., Chinwalla, V. and Harte, P. J. (1995). Trithorax is required to maintain engrailed expression in a subset of engrailed-expressing cells. Mech Dev 52: 89-98
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
Burke, T. W. and Kadonaga, J. T. (1996). Drosophila TFIID binds to a conserved downstream basal promoter element that is present in many TATA-box-deficient promoters. Genes and Development 10: 711-727
Cadigan, K. M., Grossniklaus, U. and Gehring, W. J., (1994). Localized expression of sloppy paired protein maintains the polarity of Drosophila parasegments. Genes Dev 8: 899-913
Chen, E. H. and Baker. B. S. (1997). Compartmental organization of the Drosophila genital imaginal discs. Development 124: 205-218
Condron, B. G., Patel, N. H. and Zinn, K. (1994). Engrailed controls glial/neuronal cell fate decisions at the midline of the central nervous system. Neuron 13: 541-554
Danielian, P. S., McMahon, A. P. (1996). Engrailed-1 as a target of the WNT-1 signalling pathway in the vertebrate midbrain development. Nature 383: 332-334
de Celis, J. F. and Rulz-Gomez, M. (1995). groucho and hedgehog regulate engrailed expression in the anterior compartment of the Drosophila wing. Development 121: 3467-76
DiNardo, S. and O'Farrell, P. H. (1987). Establishment and refinement of segmental pattern in the Drosophila embryo: spatial control of engrailed expression by pair-rule genes. Genes Dev. 1: 1212-25
Fauvarque, M.O., Zuber, V and Dura, J. M. (1995). Regulation of polyhomeotic transcription may involve local changes in chromatin activity in Drosophila. Mech Dev 52: 343-355
Fjose, A., McGinnis, W.J. and Gehring, W.J. (1985). Isolation of a homoeo box-containing gene from the engrailed region of Drosophila and the spacial distribution of its transcripts. Nature 313: 284-89
Florence, B., et al. (1997). Ftz-F1 is a cofactor in Ftz activation of the Drosophila engrailed gene. Development 124: 839-847
Fujioka, M., Jaynes, J. B. and Goto, T. (1995). Early even-skipped stripes act as morphogenetic gradients at the single cell level to establish engrailed expression. Development 121: 4371-4382
Goldsborough, A. S. and Kornberg, T. B. (1994). Allele-specific quantification of Drosophila engrailed and invected transcripts. Proc Natl Acad Sci 91: 12696-12700
Guillen, I., et al. (1995). The function of engrailed and the specification of Drosophila wing pattern. Development 121: 3447-3456
Han, K and Manley, J. L. (1993). Functional domains of the Drosophila Engrailed protein. EMBO J. 12: 2723-33
Hidalgo, A. and Ingham, P. (1990). Cell patterning in the Drosophila segment: spatial regulation of the segment polarity gene patched. Development 110: 291-301
Itasaki, N. and Nakamura, H. (1996). A role for gradient en expression in positional specification on the optic tectum. Neuron 16: 55-62
Kassis, J. A. (1994). Unusual properties of regulatory DNA from the Drosophila engrailed gene: three "pairing-sensitive" sites within a 1.6-kb region. Genetics 136: 1025-38
Kornberg, T., et al. (1985). The engrailed locus of Drosophila: in situ localization of transcripts reveals compartment-specific expression. Cell 40: 45-53
Kuhn, D. T., et al. (1995). Analysis of the genes involved in organizing the tail segments of the Drosophila melanogaster embryo. Mech. Dev. 53: 3-13
LaJeunesse, D. and Shearn, A. (1995). Trans-regulation of thoracic homeotic selector genes of the Antennapedia and bithorax complexes by the trithorax group genes: absent, small, and homeotic discs 1 and 2. Mech. Dev. 53: 123-39
Laufer, E., et al. (1997). Expression of Radical fringe in limb-bud ectoderm regulates apical ectodermal ridge formation. Nature 386: 366-373
Lawrence, P., Sanson, B. and Vincent, J.-P. (1996). Compartments, wingless and engrailed: patterning and the ventral epidermis of Drosophila embryos. Development 122: 4095-4103
Lee, S. M. K., et al. (1997). Evidence that FGF8 signalling from the midbrain-hindbrain junction regulates growth and polarity in the developing midbrain. Development 124: 959-969.
Lundell, M. J., et al. (1996). The engrailed and huckebein genes are essential for development of serotonin neurons in the Drosophila CNS. Mol. Cell. Neurosci. 7: 46-61
Loomis, C. A., et al. (1996). The mouse Engrailed-1 gene and ventral limb patterning. Nature 382: 360-363
Macias, A., Pelaz, S. and Morata, G., (1994). Genetic factors controlling the expression of the abdominal-A gene of Drosophila within its domain. Mech Dev 46: 15-25
Mann, R. S. (1994). Engrailed-mediated repression of Ultrabithorax is necessary for the parasegment 6 identity in Drosophila. Development 120: 3205-3212
Manoukian, A. S., and Krause, H. M. (1993). Control of segmental asymmetry in Drosophila embryos. Development 118: 785-796
McDonald, J. A. and Doe, C. Q. (1997). Establishing neuroblast-specific gene expression in the Drosophila CNS: huckebein is activated by Wingless and Hedgehog and repressed by Engrailed and Gooseberry. Development 124: 1079-1087
Mullen, J. R. and DiNardo, S. (1995). Establishing parasegments in Drosophila embryos: roles of odd-skipped and naked genes. Dev. Biol. 169: 295-308
Nambu, J. R., et al. (1990). The single-minded gene of Drosophila is required for the expression of genes important for the development of CNS midline cells. Cell 63: 63-75
Peltenburg, L. T. C. and Murre, C. (1996). Engrailed and Hox homeodomain proteins contain a related Pbx interaction motif that recognizes a common structure present in Pbx. EMBO J. 15: 3385-93
Poole, S.J. et. al. (1985). The engrailed locus of Drosophila: Structural analysis of an embryonic transcript. Cell 40: 37-43
Retaux, S., McNeill, L. and Harris, W.A. (1996). Engrailed, retinotectal targeting and axonal patterning in the midbrain during Xenopus development: an antisense study. Neuron 16: 63-75
Rodriguez-Esteban, C., et al. (1997). Radical fringe positions the apical ectodermal ridge at the dorsoventral boundary of the vertebrate limb. Nature 386: 360-366
Rogers, B. T. and Kaufman, T. C. (1996). Structure of the insect head as revealed by the EN protein pattern in developing embryos. Development 122, 3419-3432
Romani, S., et al. (1996). Krüppel, a Drosophila segmentation gene, participates in the specification of neurons and glial cells. Mech. Dev. 60: 95-107
Saenz-Robles, M. T., et al. (1995). Selection and characterization of sequences with high affinity for engrailed proteins of Drosophila. Mech. Dev. 53: 185-195
Sanicola, M., et al. (1995). Drawing a stripe in Drosophila imaginal disks: negative regulation of decapentaplegic and patched expression by engrailed. Genetics 139: 745-756
Schmidt-Ott, U. and Technau, G. M. (1992). Expression of en and wg in the embryonic head and brain of Drosophila indicates a refolded band of seven segment remnants. Development 116: 111-25
Schwartz, C., Locke, J., Nishida, C. and Kornberg,T. B. (1995). Analysis of cubitus interruptus regulation in Drosophila embryos and imaginal disks. Development 121: 1625-1635
Serrano, N., et al. (1995). polyhomeotic appears to be a target of engrailed regulation in Drosophila. Development 121: 1691
Siegfried, E. L., Chou. T. B. and Perrimon, N. (1992). wingless signaling acts through zeste-white 3, the Drosophila homolog of glycogen synthase kinase-3, to regulate engrailed and establish cell fate. Cell 71: 1167-79
Siegfried, E. L., Wilder, E. and Perrimon, N. (1994). Components of wingless signalling in Drosophila. Nature 367: 76-80
Simonds, A. J., et al. (1995). Distinguishable functions for engrailed and invected in anterior-posterior patterning in the Drosophila wing. Nature 376: 424-427
Song, D. L., et al. (1996). Two Pax-binding sites are required for early embryonic brain expression of an Engrailed-2 transgene. Development 122: 627-635
Strutt, D. I. and Mlodzik, M. (1996). The regulation of hedgehog and decapentaplegic during Drosophila eye imaginal disc development. Mech. Dev. 58: 39-50
Tabata, T. , Eaton, S. and Kornberg, T.B. (1992). The Drosophila hedgehog gene is expressed specifically in posterior compartment cells and is a target of engrailed regulation. Genes Dev. 6: 2635-2645
Tabata, T., et al. (1995). Creating a Drosophila wing de novo, the role of engrailed, and the compartment border hypothesis. Development 121: 3359-3369
van de Wetering, M., et al. (1997). Armadillo coactivates transcription driven by the product of the Drosophila segment polarity gene dTCF. Cell 88: 789-799
van Dijk, M. A. and Murre, C. (1994). Extradenticle raises the DNA binding specificity of homeotic selector gene products. Cell 78: 617-624
Vincent, J. P. and Lawrence, P. A. (1994). Drosophila wingless sustains engrailed expression only in adjoining cells: evidence from mosaic embryos. Cell 77: 909-15
Wurst, W., Auerbach, A. B. and Joyner, A. L. (1994). Multiple developmental defects in Engrailed-1 mutant mice: an early mid-hindbrain deletion and patterning defects in forelimbs and sternum. Development 120: 2065-2075
Younossi-Hartenstein, A., et al. (1996). Early neurogenesis of the Drosophila
brain. J. Comp. Neur. 370: 313-329
HOM-C
HOX-Pro