Identification of Polycomb and trithorax group responsive elements in the regulatory region of the Drosophila homeotic gene Sex combs reduced.

Genetics ◽  
1995 ◽  
Vol 139 (2) ◽  
pp. 797-814 ◽  
Author(s):  
J G Gindhart ◽  
T C Kaufman
Keyword(s):  
Regulatory Region ◽  
Polycomb Group ◽  
Homeotic Gene ◽  
Regulatory Sequences ◽  
Dependent Manner ◽  
Gene Products ◽  
Dna Fragments ◽  
Sex Combs Reduced ◽  
Trithorax Group ◽  
Sex Combs

Abstract The Drosophila homeotic gene Sex combs reduced (Scr) is necessary for the establishment and maintenance of the morphological identity of the labial and prothoracic segments. In the early embryo, its expression pattern is established through the activity of several gap and segmentation gene products, as well as other transcription factors. Once established, the Polycomb group (Pc-G) and trithorax group (trx-G) gene products maintain the spatial pattern of Scr expression for the remainder of development. We report the identification of DNA fragments in the Scr regulatory region that may be important for its regulation by Polycomb and trithorax group gene products. When DNA fragments containing these regulatory sequences are subcloned into P-element vectors containing a white minigene, transformants containing these constructs exhibit mosaic patterns of pigmentation in the adult eye, indicating that white minigene expression is repressed in a clonally heritable manner. The size of pigmented and nonpigmented clones in the adult eye suggests that the event determining whether a cell in the eye anlagen will express white occurs at least as early as the first larval instar. The amount of white minigene repression is reduced in some Polycomb group mutants, whereas repression is enhanced in flies mutant for a subset of trithorax group loci. The repressor activity of one fragment, normally located in Scr Intron 2, is increased when it is able to homologously pair, a property consistent with genetic data suggesting that Scr exhibits transvection. Another Scr regulatory fragment, normally located 40 kb upstream of the Scr promoter, silences ectopic expression of an Scr-lacZ fusion gene in the embryo and does so in a Polycomb-dependent manner. We propose that the regulatory sequences located within these DNA fragments may normally mediate the regulation of Scr by proteins encoded by members of the Polycomb and trithorax group loci.

scholarly journals Characterization of the cis-regulatory region of the Drosophila homeotic gene Sex combs reduced.

Genetics ◽  
1995 ◽  
Vol 139 (2) ◽  
pp. 781-795
Author(s):  
J G Gindhart ◽  
A N King ◽  
T C Kaufman
Keyword(s):  
Gene Expression ◽  
Reporter Gene ◽  
Regulatory Region ◽  
Reporter Gene Expression ◽  
Homeotic Gene ◽  
Regulatory Sequences ◽  
Dna Fragments ◽  
Sex Combs Reduced ◽  
Posterior Midgut ◽  
Sex Combs

Abstract The Drosophila homeotic gene Sex combs reduced (Scr) controls the segmental identity of the labial and prothoracic segments in the embryo and adult. It encodes a sequence-specific transcription factor that controls, in concert with other gene products, differentiative pathways of tissues in which Scr is expressed. During embryogenesis, Scr accumulation is observed in a discrete spatiotemporal pattern that includes the labial and prothoracic ectoderm, the subesophageal ganglion of the ventral nerve cord and the visceral mesoderm of the anterior and posterior midgut. Previous analyses have demonstrated that breakpoint mutations located in a 75-kb interval, including the Scr transcription unit and 50 kb of upstream DNA, cause Scr misexpression during development, presumably because these mutations remove Scr cis-regulatory sequences from the proximity of the Scr promoter. To gain a better understanding of the regulatory interactions necessary for the control of Scr transcription during embryogenesis, we have begun a molecular analysis of the Scr regulatory interval. DNA fragments from this 75-kb region were subcloned into P-element vectors containing either an Scr-lacZ or hsp70-lacZ fusion gene, and patterns of reporter gene expression were assayed in transgenic embryos. Several fragments appear to contain Scr regulatory sequences, as they direct reporter gene expression in patterns similar to those normally observed for Scr, whereas other DNA fragments direct Scr reporter gene expression in developmentally interesting but non-Scr-like patterns during embryogenesis. Scr expression in some tissues appears to be controlled by multiple regulatory elements that are separated, in some cases, by more than 20 kb of intervening DNA. Interestingly, regulatory sequences that direct reporter gene expression in an Scr-like pattern in the anterior and posterior midgut are imbedded in the regulatory region of the segmentation gene fushi tarazu (ftz), which is normally located between 10 and 20 kb 5' of the Scr transcription start site. This analysis provides an entry point for the study of how Scr transcription is regulated at the molecular level.

scholarly journals An Organizational Hub of Developmentally Regulated Chromatin Loops in the Drosophila Antennapedia Complex

2015 ◽  
Vol 35 (23) ◽  
pp. 4018-4029 ◽  
Author(s):  
Mo Li ◽  
Zhibo Ma ◽  
Jiayang K. Liu ◽  
Sharmila Roy ◽  
Sapna K. Patel ◽  
...  
Keyword(s):  
Gene Regulation ◽  
Regulatory Region ◽  
Regulatory Sequence ◽  
Regulatory Sequences ◽  
Developmentally Regulated ◽  
Sex Combs Reduced ◽  
Content Type ◽  
Chromatin Loops ◽  
Sex Combs ◽  
Genomic Environment

Chromatin boundary elements (CBEs) are widely distributed in the genome and mediate formation of chromatin loops, but their roles in gene regulation remain poorly understood. The complex expression pattern of theDrosophilahomeotic geneSex combs reduced(Scr) is directed by an unusually long regulatory sequence harboring diverseciselements and an intervening neighbor genefushi tarazu(ftz). Here we report the presence of a multitude of CBEs in theScrregulatory region. Selective and dynamic pairing among these CBEs mediates developmentally regulated chromatin loops. In particular, the SF1 boundary plays a central role in organizing two subsets of chromatin loops: one subset enclosesftz, limiting its access by the surroundingScrenhancers and compartmentalizing distinct histone modifications, and the other subset subdivides theScrregulatory sequences into independent enhancer access domains. We show that these CBEs exhibit diverse enhancer-blocking activities that vary in strength and tissue distribution. Tandem pairing of SF1 and SF2, two strong CBEs that flank theftzdomain, allows the distal enhancers to bypass their block in transgenicDrosophila, providing a mechanism for the endogenousScrenhancer to circumvent theftzdomain. Our study demonstrates how an endogenous CBE network, centrally orchestrated by SF1, could remodel the genomic environment to facilitate gene regulation during development.

scholarly journals DSP1, an HMG-like Protein, Is Involved in the Regulation of Homeotic Genes

Genetics ◽  
2001 ◽  
Vol 157 (1) ◽  
pp. 237-244
Author(s):  
M Decoville ◽  
E Giacomello ◽  
M Leng ◽  
D Locker
Keyword(s):  
Null Allele ◽  
Genetic Interaction ◽  
Polycomb Group ◽  
Homeotic Genes ◽  
Loss Of Function ◽  
Sex Combs Reduced ◽  
Trithorax Group ◽  
Sex Combs ◽  
Normal Expression ◽  
Sex Comb

Abstract The Drosophila dsp1 gene, which encodes an HMG-like protein, was originally identified in a screen for corepressors of Dorsal. Here we report that loss of dsp1 function causes homeotic transformations resembling those associated with loss of function in the homeotic genes Sex combs reduced (Scr), Ultrabithorax (Ubx), and Abdominal-B. The expression pattern of Scr is altered in dsp1 mutant imaginal discs, indicating that dsp1 is required for normal expression of this gene. Genetic interaction studies reveal that a null allele of dsp1 enhances trithorax-group gene (trx-G) mutations and partially suppresses Polycomb-group gene (Pc-G) mutations. On the contrary, overexpression of dsp1 induces an enhancement of the transformation of wings into halteres and of the extra sex comb phenotype of Pc. In addition, dsp1 male mutants exhibit a mild transformation of A4 into A5. Comparison of the chromatin structure at the Mcp locus in wild-type and dsp1 mutant embryos reveals that the 300-bp DNase I hypersensitive region is absent in a dsp1 mutant context. We propose that DSP1 protein is a chromatin remodeling factor, acting as a trx-G or a Pc-G protein depending on the considered function.

Transvection and Silencing of the Scr Homeotic Gene of Drosophila melanogaster

Genetics ◽  
2002 ◽  
Vol 161 (2) ◽  
pp. 733-746
Author(s):  
Jeffrey W Southworth ◽  
James A Kennison
Keyword(s):  
Drosophila Melanogaster ◽  
Chromosomal Aberrations ◽  
Regulatory Element ◽  
Regulatory Elements ◽  
Anomalous Behavior ◽  
Polycomb Group ◽  
Homeotic Gene ◽  
Polycomb Group Proteins ◽  
Sex Combs Reduced ◽  
In Cis

Abstract The Sex combs reduced (Scr) gene specifies the identities of the labial and first thoracic segments in Drosophila melanogaster. In imaginal cells, some Scr mutations allow cis-regulatory elements on one chromosome to stimulate expression of the promoter on the homolog, a phenomenon that was named transvection by Ed Lewis in 1954. Transvection at the Scr gene is blocked by rearrangements that disrupt pairing, but is zeste independent. Silencing of the Scr gene in the second and third thoracic segments, which requires the Polycomb group proteins, is disrupted by most chromosomal aberrations within the Scr gene. Some chromosomal aberrations completely derepress Scr even in the presence of normal levels of all Polycomb group proteins. On the basis of the pattern of chromosomal aberrations that disrupt Scr gene silencing, we propose a model in which two cis-regulatory elements interact to stabilize silencing of any promoter or cis-regulatory element physically between them. This model also explains the anomalous behavior of the Scx allele of the flanking homeotic gene, Antennapedia. This allele, which is associated with an insertion near the Antennapedia P1 promoter, inactivates the Antennapedia P1 and P2 promoters in cis and derepresses the Scr promoters both in cis and on the homologous chromosome.

scholarly journals Novel patterns of homeotic protein accumulation in the head of the Drosophila embryo

Development ◽  
1989 ◽  
Vol 105 (1) ◽  
pp. 167-174 ◽  
Author(s):  
J.W. Mahaffey ◽  
R.J. Diederich ◽  
T.C. Kaufman
Keyword(s):  
Nerve Cord ◽  
Protein Product ◽  
Drosophila Embryo ◽  
Protein Accumulation ◽  
Homeotic Genes ◽  
Gene Products ◽  
Ventral Region ◽  
Sex Combs Reduced ◽  
Sex Combs ◽  
Dorsal Epidermis

Antibodies that specifically recognize proteins encoded by the homeotic genes: Sex combs reduced, Deformed, labial and proboscipedia, were used to follow the distribution of these gene products during embryogenesis. The position of engrailed-expressing cells was used as a reference and staining conditions were established that could distinguish, among cells expressing engrailed, one of the homeotic proteins or both. Our observations demonstrate two important facts about establishing identity in the head segments. First, in contrast to the overlapping pattern of homeotic gene expression in the trunk segments, we observe a non-overlapping pattern in the head for those homeotic proteins required during embryogenesis. In contrast, the spatial accumulation of the protein product of the non-vital proboscipedia locus overlaps partially with the distribution of the Deformed and Sex combs reduced proteins in the maxillary and labial segments, respectively. Second, two of the proteins, Sex combs reduced and Deformed, have different dorsal and ventral patterns of accumulation. Dorsally, these proteins are expressed in segmental domains while, within the ventral region, a parasegmental register is observed. The boundary where this change in pattern occurs coincides with the junction between the ventral neurogenic region and the dorsal epidermis. After contraction of the germ band, when the nerve cord has completely separated from the epidermis, the parasegmental pattern is observed only within the ventral nerve cord while a segmental register is maintained throughout the epidermis.

Evolution of the insect body plan as revealed by the Sex combs reduced expression pattern

Development ◽  
1997 ◽  
Vol 124 (1) ◽  
pp. 149-157 ◽  
Author(s):  
B.T. Rogers ◽  
M.D. Peterson ◽  
T.C. Kaufman
Keyword(s):  
Morphological Evolution ◽  
Expression Patterns ◽  
Morphological Characters ◽  
Homeotic Gene ◽  
Homeotic Genes ◽  
Cell Fates ◽  
Sex Combs Reduced ◽  
Sex Combs ◽  
Evolutionarily Conserved ◽  
Epidermal Expression

The products of the HOM/Hox homeotic genes form a set of evolutionarily conserved transcription factors that control elaborate developmental processes and specify cell fates in many metazoans. We examined the expression of the ortholog of the homeotic gene Sex combs reduced (Scr) of Drosophila melanogaster in insects of three divergent orders: Hemiptera, Orthoptera and Thysanura. Our data reflect how the conservation and variation of Scr expression has affected the morphological evolution of insects. Whereas the anterior epidermal expression of Scr, in a small part of the posterior maxillary and all of the labial segment, is found to be in common among all four insect orders, the posterior (thoracic) expression domains vary. Unlike what is observed in flies, the Scr orthologs of other insects are not expressed broadly over the first thoracic segment, but are restricted to small patches. We show here that Scr is required for suppression of wings on the prothorax of Drosophila. Moreover, Scr expression at the dorsal base of the prothoracic limb in two other winged insects, crickets (Orthoptera) and milkweed bugs (Hemiptera), is consistent with Scr acting as a suppressor of prothoracic wings in these insects. Scr is also expressed in a small patch of cells near the basitarsal-tibial junction of milkweed bugs, precisely where a leg comb develops, suggesting that Scr promotes comb formation, as it does in Drosophila. Surprisingly, the dorsal prothoracic expression of Scr is also present in the primitively wingless firebrat (Thysanura) and the leg patch is seen in crickets, which have no comb. Mapping both gene expression patterns and morphological characters onto the insect phylogenetic tree demonstrates that in the cases of wing suppression and comb formation the appearance of expression of Scr in the prothorax apparently precedes these specific functions.

Maintenance of the active and inactive states

2019 ◽  
pp. 80-92
Author(s):  
John C. Lucchesi
Keyword(s):  
Transcriptional Repression ◽  
Homologous Chromosome ◽  
Regulatory Region ◽  
Histone H3 ◽  
Epigenetic Modifications ◽  
Polycomb Group ◽  
Histone H2a ◽  
Trithorax Group ◽  
Non Coding Rnas ◽  
Somatic Pairing

The maintenance of a gene in an active or inactive state is carried out by epigenetic modifications of the histones and of the DNA itself. Two major classes of complexes (PRC1 and PRC2), containing Polycomb group (PcG) proteins mediate transcriptional repression. PRC2 trimethylates histone H3 at lysine 27, a modification that attracts PRC1 leading to the ubiquitination of histone H2A. Variant PRC1 complexes can be targeted first, and mono-ubiquitinated histone H2A recruits PRC2 complexes that serve as the target for canonical PRC1 complexes. PRC2 can be targeted to sites of repression by associating with long non-coding RNAs. Trithorax group (TrxG) proteins form complexes that counteract PcG-mediated repression. Some subunits of these complexes maintain and enhance transcription by carrying out different lysine methylations (H3K4me, H3K36me and H3K79me) that are associated with active gene function; other subunits remodel chromatin by displacing and repositioning nucleosomes. Additional effects on transcription are transvections, whereby somatic pairing allows the regulatory region of one allele of a gene to influence the activity of the promoter of the allele on the homologous chromosome

scholarly journals Novel regulation of the homeotic gene Scr associated with a crustacean leg-to-maxilliped appendage transformation

Development ◽  
1999 ◽  
Vol 126 (6) ◽  
pp. 1121-1128 ◽  
Author(s):  
A. Abzhanov ◽  
T.C. Kaufman
Keyword(s):  
Expression Patterns ◽  
Homeotic Gene ◽  
Homeotic Genes ◽  
Porcellio Scaber ◽  
Sex Combs Reduced ◽  
Early Embryos ◽  
Sex Combs ◽  
Thoracic Legs ◽  
Post Transcriptional Regulation ◽  
Late Stages

Homeotic genes are known to be involved in patterning morphological structures along the antero-posterior axis of insects and vertebrates. Because of their important roles in development, changes in the function and expression patterns of homeotic genes may have played a major role in the evolution of different body plans. For example, it has been proposed that during the evolution of several crustacean lineages, changes in the expression patterns of the homeotic genes Ultrabithorax and abdominal-A have played a role in transformation of the anterior thoracic appendages into mouthparts termed maxillipeds. This homeotic-like transformation is recapitulated at the late stages of the direct embryonic development of the crustacean Porcellio scaber (Oniscidea, Isopoda). Interestingly, this morphological change is associated with apparent novelties both in the transcriptional and post-transcriptional regulation of the Porcellio scaber ortholog of the Drosophila homeotic gene, Sex combs reduced (Scr). Specifically, we find that Scr mRNA is present in the second maxillary segment and the first pair of thoracic legs (T1) in early embryos, whereas protein accumulates only in the second maxillae. In later stages, however, high levels of SCR appear in the T1 legs, which correlates temporally with the transformation of these appendages into maxillipeds. Our observations provide further insight into the process of the homeotic leg-to-maxilliped transformation in the evolution of crustaceans and suggest a novel regulatory mechanism for this process in this group of arthropods.

scholarly journals Distribution of the Sex combs reduced Gene Products in Drosophila melanogaster

Genetics ◽  
1987 ◽  
Vol 117 (1) ◽  
pp. 51-60
Author(s):  
James W Mahaffey ◽  
Thomas C Kaufman
Keyword(s):  
Central Nervous System ◽  
Nervous System ◽  
Temporal Distribution ◽  
Spatial And Temporal Distribution ◽  
Gene Products ◽  
Genetic Studies ◽  
Sex Combs Reduced ◽  
Sex Combs ◽  
The Central Nervous System ◽  
Differential Timing

ABSTRACT The spatial and temporal distribution of RNA and protein encoded by the homeotic Sex combs reduced (Scr) gene were examined during Drosophila development. The gene products are present in the epidermis of both the labial and first thoracic segments as would be predicted from prior genetic studies. However, the pattern in the central nervous system (CNS) and mesoderm is further restricted; the major expression located in the labial neuromere of the CNS and the mesoderm of the first thoracic segment. The spatial restriction within the CNS is correlated with and may be due to a differential timing of expression in the labial and first thoracic ectoderm. The labial ectoderm accumulates the Scr RNA prior to segregation of the neuroblasts while expression in the first thoracic ectoderm occurs after neuroblast segregation. The protein is also observed in the subesophageal ganglia of both larvae and adults, as well as in the labial and first thoracic imaginal discs. Surprisingly, the protein is also present to a lesser extent in second and third thoracic leg discs.

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