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PR00026

Identifier
ENGRAILED  [View Relations]  [View Alignment]  
Accession
PR00026
No. of Motifs
2
Creation Date
08-JUN-1993  (UPDATE 14-JUN-1999)
Title
Engrailed homeodomain signature
Database References

PROSITE; PS00033 ENGRAILED
INTERPRO; IPR000747
Literature References
1. XUE, Z.G., GEHRING, W.J. AND LE DOURAIN, N.M.
Quox-1, A quail homeobox gene expressed in the embryonic central nervous
system, including the forebrain.
PROC.NATL.ACAD.SCI.U.S.A. 88(6) 2427-2431 (1991).
 
2. GEHRING, W.J.
Homeo boxes in the study of development.
SCIENCE 236 1245-1252 (1987).
 
3. ANGERER, L.M., DOLECKI, G.J., GAGNON, M.L., LUM, R., WANG, G., YANG, Q.,
HUMPHREYS, T. AND ANGERER, R.C.
Progressively restricted expression of a homeo box gene within the aboral
ectoderm of developing sea urchin embryos.
GENES DEV. 3(3) 370-383 (1989).
 
4. SASAKI, H., YOKOYAMA, E. AND KUROIWA, A.
Specific DNA-binding of the 2 chicken deformed fmily homeodomain proteins,
Chox-1.4 AND Chox-A.
NUCLEIC ACIDS RES. 18 1739-1747 (1990).
 
5. BRENNAN, R.G., TAKEDA, Y., KIM, J., ANDERSON, W.F. AND MATTHEWS, B.W.
Crystallization of a complex of cro repressor with a 17 base pair operator.
J.MOL.BIOL. 188 115-118 (1986).

Documentation
Organisms develop according to a precise program that specifies the body
plan in intricate detail and also determines the timing of developmental
events [1]. The highly complex nature of these events has suggested that 
the process may be regulated by proteins capable of controlling the 
temporal and spatial expression of many structural genes. The genes for 
this process were first discovered as homeotic mutations in Drosophila [2],
and many similar genes are now known in a wide variety of organisms.
 
Proteins that regulate developmental gene expression are nuclear proteins 
[3] that contain a conserved domain known as the homeobox, the flanking
sequences of which differ considerably among different proteins. The homeo 
domain includes the helix-turn-helix (HTH) motif, which binds to DNA in a 
sequence-specific manner to exert a temporal and spatial regulation of 
developmental gene expression [4]. The second helix of this motif binds 
to DNA via a number of hydrogen bonds and hydrophobic interactions, which
occur between specific side chains and the exposed bases and thymine methyl
groups within the major groove. The first helix may help to stablise the 
structure [5].
 
Many homeodomain-containing proteins have now been sequenced and, while
the homeodomain-flanking regions vary, characteristic conserved sequences
upstream of the domain allow the proteins to be grouped into 3 subfamilies:
the so-called antennapedia, engrailed and `paired box' proteins. Engrailed 
plays an important role in Drosophila segmentation and neurogenesis, 
affecting genes in posterior compartments of the developing embryo. It is 
also required for the development of the central nervous system. Homologues
found in other species may play a role in neurogenesis, possibly in both 
the compartmentalisation of the developing neural tube and specification of
particular neuronal populations. Members of the engrailed subfamily of
proteins contain a conserved region of 20 amino acids located to the
C-terminal of the homeobox, the specific function of which is unclear.
 
ENGRAILED is a 2-element fingerprint that provides a signature for the
engrailed-type homeobox proteins. The fingerprint was derived from an 
initial alignment of 6 sequences: motif 2 encodes the conserved region to
the C-terminus of the homeobox (cf. PROSITE pattern ENGRAILED (PS00033)).
Two iterations on OWL20.0 were required to reach convergence, at which 
point a true set comprising 20 sequences was identified.
 
An update on SPTR37_9f identified a true set of 25 sequences.
Summary Information
25 codes involving  2 elements
Composite Feature Index
22525
12
True Positives
HME1_BRARE    HME1_CHICK    HME1_HUMAN    HME1_MOUSE    
HME2_BRARE HME2_CHICK HME2_HUMAN HME2_MOUSE
HME3_BRARE HMEC_XENLA HMED_XENLA HMEN_ANOGA
HMEN_ARTSF HMEN_BOMMO HMEN_DROME HMEN_DROVI
HMIN_BOMMO HMIN_DROME HX11_HUMAN HX11_MOUSE
O76848 P90688 Q25212 Q26371
Q26601
Sequence Titles
HME1_BRARE  HOMEOBOX PROTEIN ENGRAILED-1 - BRACHYDANIO RERIO (ZEBRAFISH) (ZEBRA DANIO). 
HME1_CHICK HOMEOBOX PROTEIN ENGRAILED-1 (GG-EN-1) - GALLUS GALLUS (CHICKEN).
HME1_HUMAN HOMEOBOX PROTEIN ENGRAILED-1 (HU-EN-1) - HOMO SAPIENS (HUMAN).
HME1_MOUSE HOMEOBOX PROTEIN ENGRAILED-1 (MO-EN-1) - MUS MUSCULUS (MOUSE).
HME2_BRARE HOMEOBOX PROTEIN ENGRAILED-2 (ZF-EN-2) - BRACHYDANIO RERIO (ZEBRAFISH) (ZEBRA DANIO).
HME2_CHICK HOMEOBOX PROTEIN ENGRAILED-2 (GG-EN-2) - GALLUS GALLUS (CHICKEN).
HME2_HUMAN HOMEOBOX PROTEIN ENGRAILED-2 (HU-EN-2) - HOMO SAPIENS (HUMAN).
HME2_MOUSE HOMEOBOX PROTEIN ENGRAILED-2 (MO-EN-2) - MUS MUSCULUS (MOUSE).
HME3_BRARE HOMEOBOX PROTEIN ENGRAILED-3 (ZF-EN-1) - BRACHYDANIO RERIO (ZEBRAFISH) (ZEBRA DANIO).
HMEC_XENLA HOMEOBOX PROTEIN ENGRAILED-2A (EN-2A) (EN2 1.4) - XENOPUS LAEVIS (AFRICAN CLAWED FROG).
HMED_XENLA HOMEOBOX PROTEIN ENGRAILED-2B (EN-2B) (EN2 MABEN) - XENOPUS LAEVIS (AFRICAN CLAWED FROG).
HMEN_ANOGA SEGMENTATION POLARITY HOMEOBOX PROTEIN ENGRAILED - ANOPHELES GAMBIAE (AFRICAN MALARIA MOSQUITO).
HMEN_ARTSF HOMEOBOX PROTEIN ENGRAILED - ARTEMIA SANFRANCISCANA (BRINE SHRIMP) (ARTEMIA FRANCISCANA).
HMEN_BOMMO SEGMENTATION POLARITY HOMEOBOX PROTEIN ENGRAILED - BOMBYX MORI (SILK MOTH).
HMEN_DROME SEGMENTATION POLARITY HOMEOBOX PROTEIN ENGRAILED - DROSOPHILA MELANOGASTER (FRUIT FLY).
HMEN_DROVI SEGMENTATION POLARITY HOMEOBOX PROTEIN ENGRAILED - DROSOPHILA VIRILIS (FRUIT FLY).
HMIN_BOMMO HOMEOBOX PROTEIN INVECTED - BOMBYX MORI (SILK MOTH).
HMIN_DROME HOMEOBOX PROTEIN INVECTED - DROSOPHILA MELANOGASTER (FRUIT FLY).
HX11_HUMAN HOMEOBOX PROTEIN HOX-11 (TCL-3 PROTO-ONCOGENE) - HOMO SAPIENS (HUMAN).
HX11_MOUSE HOMEOBOX PROTEIN HOX-11 (T-CELL LEUKEMIA HOMEOBOX 1) (HOMEOBOX TLX-1) - MUS MUSCULUS (MOUSE).
O76848 HOMEOBOX PROTEIN - CUPIENNIUS SALEI.
P90688 ENGRAILED PROTEIN - BRANCHIOSTOMA FLORIDAE (FLORIDA LANCELET) (AMPHIOXUS).
Q25212 INVECTED HOMEODOMAIN PROTEIN - JUNONIA COENIA (PEACOCK BUTTERFLY) (PRECIS COENIA).
Q26371 ENGRAILED HOMOLOG - TRIBOLIUM CASTANEUM (RED FLOUR BEETLE).
Q26601 HOMEOBOX PROTEIN ENGRAILED-LIKE SMOX-2 - SCHISTOSOMA MANSONI (BLOOD FLUKE).
Scan History
OWL20_0    2  100  NSINGLE    
OWL26_0 2 300 NSINGLE
SPTR37_9f 2 300 NSINGLE
Initial Motifs
Motif 1  width=18
Element Seqn Id St Int Rpt
EEKRPRTAFSAEQLARLK HME3_APIME 18 18 -
EDKRPRTAFSGTQLARLK HMIN_DROMO 470 470 -
DEKRPRTAFSASQLQRLK HMEN_TRIGR 36 36 -
EDKRPRTAFTAEQLQRLK HME1_MOUSE 34 34 -
EEKRPRTAFSGAQLARLK HMEN_BOMMO 279 279 -
DEKRPRTAFSGPQLARLK HMIN_BOMMO 371 371 -

Motif 2 width=18
Element Seqn Id St Int Rpt
KNGLALHLMAQGLYNHST HME1_MOUSE 97 45 -
RNPLALQLMAQGLYNHST HMEN_BOMMO 342 45 -
RNPLALQLMAQGLYNHST HMIN_BOMMO 434 45 -
KNPLALQLMAQGLYNHST HME3_APIME 81 45 -
KNPLALQLMAQGLYNHST HMIN_DROMO 533 45 -
KNDLARQLMAQGLYNHST HMEN_TRIGR 99 45 -
Final Motifs
Motif 1  width=18
Element Seqn Id St Int Rpt
EDKRPRTAFTAEQLQRLK HME1_CHICK 243 243 -
EDKRPRTAFTAEQLQRLK HME1_MOUSE 34 34 -
EDKRPRTAFTAEQLQRLK HME1_HUMAN 301 301 -
EDKRPRTAFTADQLQRLK HMEC_XENLA 175 175 -
EDKRPRTAFTAEQLQRLK HME2_CHICK 198 198 -
EDKRPRTAFTAEQLQRLK HME2_HUMAN 242 242 -
EDKRPRTAFTAEQLQRLK HME2_MOUSE 234 234 -
EDKRPRTAFTAEQLQRLK HME2_BRARE 175 175 -
EDKRPRTAFTAEQLQRLK HMED_XENLA 175 175 -
DEKRPRTAFTAEQLSRLK HMEN_ARTSF 248 248 -
EEKRPRTAFTSEQLQRLK P90688 158 158 -
DDKRPRTAFTAEQLQRLK HME1_BRARE 142 142 -
EDKRPRTAFTAEQLQRLK HME3_BRARE 171 171 -
EDKRPRTAFSGTQLARLK HMIN_DROME 470 470 -
EEKRPRTAFSNAQLQRLK HMEN_ANOGA 497 497 -
EEKRPRTAFSGAQLARLK Q26371 227 227 -
EEKRPRTAFSGAQLARLK HMEN_BOMMO 279 279 -
DEKRPRTAFSSEQLARLK HMEN_DROVI 485 485 -
DEKRPRTAFSSEQLARLK HMEN_DROME 453 453 -
DEKRPRTAFSGPQLARLK HMIN_BOMMO 371 371 -
DEKRPRTAFSGPQLARLK Q25212 38 38 -
DDKRPRTAFTADQLSRLK O76848 145 145 -
NLKRPRTSFTVPQLKRLS Q26601 422 422 -
KKKKPRTSFTRLQICELE HX11_MOUSE 202 202 -
KKKKPRTSFTRLQICELE HX11_HUMAN 200 200 -

Motif 2 width=18
Element Seqn Id St Int Rpt
KNGLALHLMAQGLYNHST HME1_CHICK 306 45 -
KNGLALHLMAQGLYNHST HME1_MOUSE 97 45 -
KNGLALHLMAQGLYNHST HME1_HUMAN 364 45 -
KNSLALHLMAQGLYNHST HMEC_XENLA 238 45 -
KNSLAVHLMAQGLYNHST HME2_CHICK 261 45 -
KNTLAVHLMAQGLYNHST HME2_HUMAN 305 45 -
KNTLAVHLMAQGLYNHST HME2_MOUSE 297 45 -
KNGLAIHLMAQGLYNHST HME2_BRARE 238 45 -
KNSLALHLMAQGLYNHAT HMED_XENLA 238 45 -
KNPLALQLMAQGLYNHST HMEN_ARTSF 311 45 -
RNGLALHLMAQGLYNHST P90688 221 45 -
KNALAMQLMAQGLYNHST HME1_BRARE 205 45 -
KNTLAVHLMAQGLYNHAT HME3_BRARE 234 45 -
KNPLALQLMAQGLYNHST HMIN_DROME 533 45 -
KNPLALQLMAQGLYNHST HMEN_ANOGA 560 45 -
KNPLALQLMAQGLYNHST Q26371 290 45 -
RNPLALQLMAQGLYNHST HMEN_BOMMO 342 45 -
KNPLALQLMAQGLYNHTT HMEN_DROVI 548 45 -
KNPLALQLMAQGLYNHTT HMEN_DROME 516 45 -
RNPLALQLMAQGLYNHST HMIN_BOMMO 434 45 -
RNPLALQLMAQGLYNHST Q25212 101 45 -
RSALALQLMAQGLYNHST O76848 208 45 -
QNCLALHLMAEGLYNHSV Q26601 485 45 -
QKSLAQPLPADPLCVHNS HX11_MOUSE 286 66 -
QKSLAQPLPADPLCVHNS HX11_HUMAN 284 66 -