Gene Symbol: Scr
Description: Sex combs reduced
Alias: BG:DS07876.2, CG1030, DmScr, Dmel\CG1030, Msc, SCR, l(3)84Af, scr, Sex combs reduced, CG1030-PA, CG1030-PB, Scr-PA, Scr-PB, multi sex combs, multiple sex comb, sex combs reduced, sex-combs reduced
Species: fruit fly
Products:     Scr

Top Publications

  1. Kosman D, Mizutani C, Lemons D, Cox W, McGinnis W, Bier E. Multiplex detection of RNA expression in Drosophila embryos. Science. 2004;305:846 pubmed
  2. Enright A, John B, Gaul U, Tuschl T, Sander C, Marks D. MicroRNA targets in Drosophila. Genome Biol. 2003;5:R1 pubmed
    ..The target prediction algorithm is freely available and can be applied to whole genome sequences using identified miRNA sequences. ..
  3. Joshi R, Sun L, Mann R. Dissecting the functional specificities of two Hox proteins. Genes Dev. 2010;24:1533-45 pubmed publisher
    ..For the Drosophila Hox protein Sex combs reduced (Scr), Exd has been shown to position a normally unstructured portion of Scr so that two basic amino acid ..
  4. Li X, Murre C, McGinnis W. Activity regulation of a Hox protein and a role for the homeodomain in inhibiting transcriptional activation. EMBO J. 1999;18:198-211 pubmed
    ..The regulation of Hox transcriptional activation functions may underlie the different functional specificities of proteins belonging to this developmental patterning family. ..
  5. Reuter R, Panganiban G, Hoffmann F, Scott M. Homeotic genes regulate the spatial expression of putative growth factors in the visceral mesoderm of Drosophila embryos. Development. 1990;110:1031-40 pubmed
    ..The two growth factor genes therefore are excellent candidates for target genes that are directly regulated by the homeotic genes. ..
  6. Percival Smith A, Teft W, Barta J. Tarsus determination in Drosophila melanogaster. Genome. 2005;48:712-21 pubmed
    ..whether Antennapedia (ANTP) cell autonomously or noncell autonomously determines tarsus identity and whether Sex combs reduced (SCR) is the HOX protein required for normal tarsus determination...
  7. Casares F, Mann R. Control of antennal versus leg development in Drosophila. Nature. 1998;392:723-6 pubmed
    ..Taken together, these data indicate that hth is an antennal selector gene, and that Antp promotes leg development by repressing hth and consequently nuclear Exd. ..
  8. Mann R, Chan S. Extra specificity from extradenticle: the partnership between HOX and PBX/EXD homeodomain proteins. Trends Genet. 1996;12:258-62 pubmed
  9. Prince F, Katsuyama T, Oshima Y, Plaza S, Resendez Perez D, Berry M, et al. The YPWM motif links Antennapedia to the basal transcriptional machinery. Development. 2008;135:1669-79 pubmed publisher
    ..BIP2 is a TATA-binding protein associated factor (also known as dTAFII3) that links ANTP to the basal transcriptional machinery. ..

More Information


  1. Papadopoulos D, Vukojevic V, Adachi Y, Terenius L, Rigler R, Gehring W. Function and specificity of synthetic Hox transcription factors in vivo. Proc Natl Acad Sci U S A. 2010;107:4087-92 pubmed publisher
    ..Here we show that synthetic Sex combs reduced (Scr) genes that encode the Scr C terminus containing the homedomain (HD) and YPWM motif (Scr-HD) are ..
  2. Heuer J, Kaufman T. Homeotic genes have specific functional roles in the establishment of the Drosophila embryonic peripheral nervous system. Development. 1992;115:35-47 pubmed
    ..The PNS was examined in embryos homozygous for mutations in the homeotic genes Sex combs reduced (Scr), Antennapedia (Antp), Ultrabithorax (Ubx), abdominal-A (abd-A) and Abdominal-B (Abd-B) with antibodies ..
  3. Uhl J, Cook T, Gebelein B. Comparing anterior and posterior Hox complex formation reveals guidelines for predicting cis-regulatory elements. Dev Biol. 2010;343:154-66 pubmed publisher
    ..Based upon these findings, we suggest a general set of guidelines to serve as a basis for designing bioinformatics algorithms aimed at identifying Hox regulatory elements using the wealth of recently sequenced genomes. ..
  4. Tayyab I, Hallahan H, Percival Smith A. Analysis of Drosophila proboscipedia mutant alleles. Genome. 2004;47:600-9 pubmed
    ..missense alleles suggests that the PB HD is required for maxillary palp and proboscis development and pb - Sex combs reduced (Scr) genetic interaction...
  5. Pattatucci A, Kaufman T. The homeotic gene Sex combs reduced of Drosophila melanogaster is differentially regulated in the embryonic and imaginal stages of development. Genetics. 1991;129:443-61 pubmed
    The Sex combs reduced (Scr) locus is unique among the genes contained within the Antennapedia complex (ANT-C) of Drosophila melanogaster in that it directs functions that are required for both cephalic and thoracic development in the ..
  6. Panzer S, Weigel D, Beckendorf S. Organogenesis in Drosophila melanogaster: embryonic salivary gland determination is controlled by homeotic and dorsoventral patterning genes. Development. 1992;114:49-57 pubmed
    ..epithelial cells derived from parasegment 2, is established by the positive action of the homeotic gene Sex combs reduced (Scr)...
  7. Brennecke J, Stark A, Russell R, Cohen S. Principles of microRNA-target recognition. PLoS Biol. 2005;3:e85 pubmed
  8. Rappailles A, Decoville M, Locker D. DSP1, a Drosophila HMG protein, is involved in spatiotemporal expression of the homoeotic gene Sex combs reduced. Biol Cell. 2005;97:779-85 pubmed
    ..The homoeotic gene Scr (Sex combs reduced) contained in the Antennapedia complex specifies segmental identity of the labial and prothoracic segments ..
  9. Bejarano F, Busturia A. Function of the Trithorax-like gene during Drosophila development. Dev Biol. 2004;268:327-41 pubmed
    ..We also suggest that GAGA factor may not have a dual activator/repressor function. Rather, Trithorax-like mutations may produce dual loss of activation and loss of repression effects. ..
  10. Bradley P, Myat M, Comeaux C, Andrew D. Posterior migration of the salivary gland requires an intact visceral mesoderm and integrin function. Dev Biol. 2003;257:249-62 pubmed
    ..These findings suggest that salivary tube dimensions may be an intrinsic property of salivary gland cells. ..
  11. Zhao J, Lazzarini R, Pick L. The mouse Hox-1.3 gene is functionally equivalent to the Drosophila Sex combs reduced gene. Genes Dev. 1993;7:343-54 pubmed
    To test whether the mouse Hox-1.3 gene is a cognate of the Drosophila Sex combs reduced (Scr) gene, we inserted a hsp 70-Hox-1.3 fusion gene into the Drosophila genome...
  12. Rieckhof G, Casares F, Ryoo H, Abu Shaar M, Mann R. Nuclear translocation of extradenticle requires homothorax, which encodes an extradenticle-related homeodomain protein. Cell. 1997;91:171-83 pubmed
    ..These data suggest a novel and evolutionarily conserved mechanism for regulating HOX activity in which a direct protein-protein interaction between EXD and HTH results in EXD's nuclear translocation. ..
  13. Beisel C, Buness A, Roustan Espinosa I, Koch B, Schmitt S, Haas S, et al. Comparing active and repressed expression states of genes controlled by the Polycomb/Trithorax group proteins. Proc Natl Acad Sci U S A. 2007;104:16615-20 pubmed
    ..Besides its presumptive function in recruiting PcG complexes to their site of action, our results now uncover that Pho plays an additional role in the repression of already induced genes. ..
  14. Ebner A, Cabernard C, Affolter M, Merabet S. Recognition of distinct target sites by a unique Labial/Extradenticle/Homothorax complex. Development. 2005;132:1591-600 pubmed
    ..Thus Lab, and most probably other Hox proteins, selects different DNA sequences in regulating downstream target genes. These observations have implications with regard to the current model of Hox-binding selectivity. ..
  15. Lelli K, Noro B, Mann R. Variable motif utilization in homeotic selector (Hox)-cofactor complex formation controls specificity. Proc Natl Acad Sci U S A. 2011;108:21122-7 pubmed publisher
    ..We find that Sex combs reduced (Scr) uses a simple interaction mechanism, where a single tryptophan-containing motif is necessary for Exd-..
  16. Slattery M, Riley T, Liu P, Abe N, Gomez Alcala P, Dror I, et al. Cofactor binding evokes latent differences in DNA binding specificity between Hox proteins. Cell. 2011;147:1270-82 pubmed publisher
    ..Together, these data suggest that emergent DNA recognition properties revealed by interactions with cofactors contribute to transcription factor specificities in vivo. ..
  17. Gindhart J, King A, Kaufman T. Characterization of the cis-regulatory region of the Drosophila homeotic gene Sex combs reduced. Genetics. 1995;139:781-95 pubmed
    The Drosophila homeotic gene Sex combs reduced (Scr) controls the segmental identity of the labial and prothoracic segments in the embryo and adult...
  18. Decoville M, Giacomello E, Leng M, Locker D. DSP1, an HMG-like protein, is involved in the regulation of homeotic genes. Genetics. 2001;157:237-44 pubmed
    ..causes homeotic transformations resembling those associated with loss of function in the homeotic genes Sex combs reduced (Scr), Ultrabithorax (Ubx), and Abdominal-B...
  19. Mahaffey J, Kaufman T. Distribution of the Sex combs reduced gene products in Drosophila melanogaster. Genetics. 1987;117:51-60 pubmed
    The spatial and temporal distribution of RNA and protein encoded by the homeotic Sex combs reduced (Scr) gene were examined during Drosophila development...
  20. Duboule D. The rise and fall of Hox gene clusters. Development. 2007;134:2549-60 pubmed
    ..By integrating these parameters, general conclusions emerge that can help solve the aforementioned dilemma. ..
  21. Andrew D, Horner M, Petitt M, Smolik S, Scott M. Setting limits on homeotic gene function: restraint of Sex combs reduced activity by teashirt and other homeotic genes. EMBO J. 1994;13:1132-44 pubmed
    ..We have identified genes that limit where the homeotic gene Sex combs reduced (Scr) can affect cell fates in the Drosophila embryo...
  22. Gambetta M, Oktaba K, Muller J. Essential role of the glycosyltransferase sxc/Ogt in polycomb repression. Science. 2009;325:93-6 pubmed publisher
    ..Polycomb repression appears to be a critical function of Sxc/Ogt in Drosophila and may be mediated by the glycosylation of Polyhomeotic. ..
  23. Scheuermann J, de Ayala Alonso A, Oktaba K, Ly Hartig N, McGinty R, Fraterman S, et al. Histone H2A deubiquitinase activity of the Polycomb repressive complex PR-DUB. Nature. 2010;465:243-7 pubmed publisher
    ..Polycomb gene silencing may thus entail a dynamic balance between H2A ubiquitination by PRC1 and dRAF, and H2A deubiquitination by PR-DUB. ..
  24. Pattatucci A, Otteson D, Kaufman T. A functional and structural analysis of the Sex combs reduced locus of Drosophila melanogaster. Genetics. 1991;129:423-41 pubmed
    We have undertaken a developmental genetic analysis of the homeotic gene Sex combs reduced (Scr) of Drosophila melanogaster by examining embryonic and adult phenotypes of mutations affecting Scr gene function...
  25. Papadopoulos D, Resendez Perez D, Cárdenas Chávez D, Villanueva Segura K, Canales del Castillo R, Felix D, et al. Functional synthetic Antennapedia genes and the dual roles of YPWM motif and linker size in transcriptional activation and repression. Proc Natl Acad Sci U S A. 2011;108:11959-64 pubmed publisher
  26. McCormick A, Core N, Kerridge S, Scott M. Homeotic response elements are tightly linked to tissue-specific elements in a transcriptional enhancer of the teashirt gene. Development. 1995;121:2799-812 pubmed
    ..We propose that local interactions between homeotic proteins and other factors effect activation of targets in proper cell types. ..
  27. Papadopoulos D, Skouloudaki K, Adachi Y, Samakovlis C, Gehring W. Dimer formation via the homeodomain is required for function and specificity of Sex combs reduced in Drosophila. Dev Biol. 2012;367:78-89 pubmed publisher
    ..We have unexpectedly found that the Drosophila Sex combs reduced (Scr) protein dimerizes in vivo via the homeodomain, whereas its closest relative, Antennapedia (Antp), does ..
  28. Joshi R, Passner J, Rohs R, Jain R, Sosinsky A, Crickmore M, et al. Functional specificity of a Hox protein mediated by the recognition of minor groove structure. Cell. 2007;131:530-43 pubmed
    ..Our results suggest that these residues, which are conserved in a paralog-specific manner, confer specificity by recognizing a sequence-dependent DNA structure instead of directly reading a specific DNA sequence. ..
  29. Noro B, Lelli K, Sun L, Mann R. Competition for cofactor-dependent DNA binding underlies Hox phenotypic suppression. Genes Dev. 2011;25:2327-32 pubmed publisher
    ..Together, these results suggest that Hox-specific motifs endow posterior Hox proteins with the ability to dominate over more anterior ones via a cofactor-dependent DNA-binding mechanism. ..
  30. Stark A, Kheradpour P, Parts L, Brennecke J, Hodges E, Hannon G, et al. Systematic discovery and characterization of fly microRNAs using 12 Drosophila genomes. Genome Res. 2007;17:1865-79 pubmed
    ..For mir-10 in particular, both arms show abundant processing, and both show highly conserved target sites in Hox genes, suggesting a possible cooperation of the two arms, and their role as a master Hox regulator. ..
  31. Percival Smith A, Weber J, Gilfoyle E, Wilson P. Genetic characterization of the role of the two HOX proteins, Proboscipedia and Sex Combs Reduced, in determination of adult antennal, tarsal, maxillary palp and proboscis identities in Drosophila melanogaster. Development. 1997;124:5049-62 pubmed
    Both Proboscipedia (PB) and Sex Combs Reduced (SCR) activities are required for determination of proboscis identity. Here we show that simultaneous removal of PB and SCR activity results in a proboscis-to-antenna transformation...
  32. Barmina O, Kopp A. Sex-specific expression of a HOX gene associated with rapid morphological evolution. Dev Biol. 2007;311:277-86 pubmed
    ..Here, we show that sex-specific regulation of the HOX gene Sex combs reduced (Scr) is associated with the origin and evolution of the Drosophila sex comb - a novel and rapidly ..
  33. Pengelly A, Copur Ö, Jackle H, Herzig A, Muller J. A histone mutant reproduces the phenotype caused by loss of histone-modifying factor Polycomb. Science. 2013;339:698-9 pubmed publisher
    ..Taken together, these analyses demonstrate that H3-K27 is the crucial physiological substrate that PRC2 modifies for Polycomb repression. ..
  34. Herz H, Madden L, Chen Z, Bolduc C, Buff E, Gupta R, et al. The H3K27me3 demethylase dUTX is a suppressor of Notch- and Rb-dependent tumors in Drosophila. Mol Cell Biol. 2010;30:2485-97 pubmed publisher
    ..In summary, these data suggest that dUTX is a suppressor of Notch- and Rbf-dependent tumors in Drosophila melanogaster and may provide a model for UTX-dependent tumorigenesis in humans. ..
  35. Tour E, Hittinger C, McGinnis W. Evolutionarily conserved domains required for activation and repression functions of the Drosophila Hox protein Ultrabithorax. Development. 2005;132:5271-81 pubmed the N termini of many Hox proteins, and is conserved in the activation domain of another Hox protein, Sex combs reduced. These results suggest that the N-terminal region containing the SSYF motif has been conserved in many Hox ..
  36. Randsholt N, Santamaria P. How Drosophila change their combs: the Hox gene Sex combs reduced and sex comb variation among Sophophora species. Evol Dev. 2008;10:121-33 pubmed publisher
    ..All contribute to spatial regulation of the Hox gene Sex combs reduced (Scr), which is crucial for comb formation...
  37. Hunter C, Kenyon C. Specification of anteroposterior cell fates in Caenorhabditis elegans by Drosophila Hox proteins. Nature. 1995;377:229-32 pubmed
    ..Together these findings suggest that factors important for target recognition by specific Hox proteins have been conserved throughout much of the animal kingdom. ..
  38. Isaac D, Andrew D. Tubulogenesis in Drosophila: a requirement for the trachealess gene product. Genes Dev. 1996;10:103-17 pubmed
    ..trh expression in the salivary duct is controlled by the homeotic gene, Sex combs reduced (Scr), and by another salivary gland gene, fork head (fkh)...
  39. Pederson J, Kiehart D, Mahaffey J. The role of HOM-C genes in segmental transformations: reexamination of the Drosophila Sex combs reduced embryonic phenotype. Dev Biol. 1996;180:131-42 pubmed
    ..Complex of Drosophila specify identity of the posterior head segments; the labial segment requires Sex combs reduced (Scr) for proper development, Deformed (Dfd) specifies maxillary and mandibular identity, and labial is ..
  40. Berry M, Gehring W. Phosphorylation status of the SCR homeodomain determines its functional activity: essential role for protein phosphatase 2A,B'. EMBO J. 2000;19:2946-57 pubmed
    b>Sex combs reduced (SCR) is a Drosophila Hox protein that determines the identity of the labial and prothoracic segments...
  41. Delorenzi M, Bienz M. Expression of Abdominal-B homeoproteins in Drosophila embryos. Development. 1990;108:323-9 pubmed
    ..Moreover, an antibody-positive Abd-B mutant suggests that these differ, in the case of m protein expression, to some extent in individual germ layers. ..
  42. Southworth J, Kennison J. Transvection and silencing of the Scr homeotic gene of Drosophila melanogaster. Genetics. 2002;161:733-46 pubmed
    The Sex combs reduced (Scr) gene specifies the identities of the labial and first thoracic segments in Drosophila melanogaster...
  43. Mann R. Why are Hox genes clustered?. Bioessays. 1997;19:661-4 pubmed
    ..Moreover, characterizing the activity of one of these mouse enhancers in Drosophila illustrates that despite many similarities, not all Hox clusters are built in the same way. ..
  44. Henderson K, Andrew D. Regulation and function of Scr, exd, and hth in the Drosophila salivary gland. Dev Biol. 2000;217:362-74 pubmed
    Salivary gland formation in the Drosophila embryo is dependent on the homeotic gene Sex combs reduced (Scr)...
  45. McGinnis W, Krumlauf R. Homeobox genes and axial patterning. Cell. 1992;68:283-302 pubmed
  46. Nelson W. Tube morphogenesis: closure, but many openings remain. Trends Cell Biol. 2003;13:615-21 pubmed
    ..This review provides a summary of these pathways based upon articles published in the Tube Morphogenesis Series in Trends in Cell Biology. ..
  47. Kennison J, Tamkun J. Dosage-dependent modifiers of polycomb and antennapedia mutations in Drosophila. Proc Natl Acad Sci U S A. 1988;85:8136-40 pubmed
    ..e., Sex combs reduced (Scr), Brista (Ba), trithorax (trx), Polycomb (Pc), Polycomblike (Pcl), and Sex comb on midleg (Scm)]...
  48. Struhl G. Genes controlling segmental specification in the Drosophila thorax. Proc Natl Acad Sci U S A. 1982;79:7380-4 pubmed
    The roles of three homeotic genes, Ubx+, Scr+, and Antp+, in the Drosophila thorax have been studied by determining the cellular phenotypes of mutations resulting in loss of gene function...
  49. Sivanantharajah L, Percival Smith A. Analysis of the sequence and phenotype of Drosophila Sex combs reduced alleles reveals potential functions of conserved protein motifs of the Sex combs reduced protein. Genetics. 2009;182:191-203 pubmed publisher
    The Drosophila Hox gene, Sex combs reduced (Scr), is required for patterning the larval and adult, labial and prothoracic segments. Fifteen Scr alleles were sequenced and the phenotypes analyzed in detail...
  50. Passner J, Ryoo H, Shen L, Mann R, Aggarwal A. Structure of a DNA-bound Ultrabithorax-Extradenticle homeodomain complex. Nature. 1999;397:714-9 pubmed
    ..Together, these protein-DNA and protein-protein interactions define the general principles by which homeotic proteins interact with Extradenticle (or Pbx1) to affect development along the anterior-posterior axis of animals. ..
  51. Roy S, Shashidhara L, Vijayraghavan K. Muscles in the Drosophila second thoracic segment are patterned independently of autonomous homeotic gene function. Curr Biol. 1997;7:222-7 pubmed
    ..We found that, unlike the mesoderm of the first and third thoracic segments which express Sex combs reduced and Antennapedia (Antp), respectively, the mesoderm of the second thoracic segment does not express any ..
  52. Andrew D, Baig A, Bhanot P, Smolik S, Henderson K. The Drosophila dCREB-A gene is required for dorsal/ventral patterning of the larval cuticle. Development. 1997;124:181-93 pubmed
    ..The lateralizing phenotype of dCREB-A mutants reveals a much broader function for CREB proteins than previously thought. ..
  53. Pare A, Lemons D, Kosman D, Beaver W, Freund Y, McGinnis W. Visualization of individual Scr mRNAs during Drosophila embryogenesis yields evidence for transcriptional bursting. Curr Biol. 2009;19:2037-42 pubmed publisher
    ..We use this method to characterize transcription at the endogenous locus of the Hox gene Sex combs reduced (Scr), by comparing a stably expressing group of cells to a group that only transiently expresses the gene...
  54. Ryoo H, Mann R. The control of trunk Hox specificity and activity by Extradenticle. Genes Dev. 1999;13:1704-16 pubmed
    ..a 37-bp element (fkh[250]) derived from the fork head (fkh) gene, a natural target of the Hox gene Sex combs reduced (Scr)...
  55. Seshaiah P, Andrew D. WRS-85D: A tryptophanyl-tRNA synthetase expressed to high levels in the developing Drosophila salivary gland. Mol Biol Cell. 1999;10:1595-608 pubmed
    ..WRS-85D expression is dependent on the homeotic gene Sex combs reduced (Scr)...
  56. Shroff S, Joshi M, Orenic T. Differential Delta expression underlies the diversity of sensory organ patterns among the legs of the Drosophila adult. Mech Dev. 2007;124:43-58 pubmed
    ..Our findings suggest that the Hox genes, Sex combs reduced (Scr) and Ultrabithorax (Ubx), establish differential expression of the proneural gene achaete (ac) by ..