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Dmel\gypsy
| General Information | |||
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| Symbol | Dmel\gypsy | Species | D.melanogaster |
| Name | gypsy element | FlyBase ID | FBte0000021 |
| Feature type | natural transposable element | ||
Recent Updates
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| Description |
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| FB2013_03 | |||
| FB2013_02 | |||
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Sequences & Components
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| Complete element (bp) |
7469
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| Terminal repeat (bp) |
482
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| Reference sequence | transposon_sequence_set.embl.txt.gz | ||
| Component genes | |||
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Sequence Accessions
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Sequence Ontology (SO)
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| Transposon type |
(Kaminker et al., 2002, Walser et al., 2006, Mugnier et al., 2008, Kulkarni and Arnosti, 2005, Kravchenko et al., 2005, Savitskaya et al., 2006, Golovnin et al., 2007, Parnell et al., 2006, Melnikova et al., 2008, Chetverina et al., 2008, Melnikova et al., 2008, Terzian et al., 2001, Llorens et al., 2008, Salenko et al., 2008, Pelisson et al., 2007)
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Insertions & Copy Number
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| Copy number and comments |
2 in euchromatin of Release 3 genome annotation, of which 1 is full length.
10 (Bayev et al.; Freund and Meselson, 1984).
TE copies retrieved from release 5.1 of the D. melanogaster genome.:11
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| Target Site Duplication | |||
| Size (bp) | |||
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Orthologs
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| Curated drosophilid orthologs | |||
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Comments
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The single full length gypsy in the sequenced strain is most likely active.
An insertion of P{SUPor-P} into a gypsy has been identified by virtue of decreasing transmission of Dp(1;f)J21A to offspring to 20%.
gypsy does not rely on germline expression for its mobilisation. The expression of a gypsy provirus in the maternal soma is necessary and sufficient for its subsequent integration into the germline of the progeny.
This soma towards germline transfer appears to be independent of the expression of the gypsy\env gene.
Cell lines carrying the permissive flam1 allele accumulate many nuclear virus-like particles, cytoplasmic dense particles and cisternae filled with fibrous material.
Two of three such cell lines have an increased copy number of the gypsy element.
The expression of gypsy encoded proteins is analysed to explore how gypsy is transmitted between generations. Assembly of gypsy particles is visualised in the follicle cells of flam females by electron microscopy, these observations provide the basis for a novel model to explain how gypsy is transmitted from generation to generation. gypsy virions appear to move through the perivitelline space during a brief developmental window and infect the oocyte, providing
a mechanism to explain gypsy insertion in the next generation.
Used in an investigation to address the relationship between retrotransposons and retroviruses and the coadaptation of these
retroelements to their host genomes. Results indicate retrotransposons are heterogeneous in contrast to retroviruses, suggesting
different modes of evolution by slippage-like mechanisms.
The nucleotide sequence of Dsub\gypsy is determined. A comparative analysis of the sequence and molecular structures of Dsub\gypsy, Dvir\gypsy and gypsy reveals gypsy is the only infectious particle, the other two have lost the ability. gypsy elements are found in D.subobscura natural populations suggesting the populations were invaded by infectious gypsy elements.
Dsub\gypsy, Dvir\gypsy and gypsy show many structural similarities including sequences necessary for transcription and regulatory and coding sequences which
suggest a common mechanism of expression. The ORF3 of Dsub\gypsy and Dvir\gypsy lack some motifs essential for the function of the "env-like" protein.
gypsy can be transmitted from flam strains in which it transposes to strains devoid of functional elements by egg plasm transfer or growing "empty" larvae in
the presence of homogenized pupae of the flam stock. Transposition of gypsy occurs only in the progeny of females homozygous for permissive alleles of flam, where gypsy transcripts are restricted to the somatic follicle cells in the ovaries. Infectious particles which then infect the oocyte
are apparently produced in these cells.
The distribution of gypsy elements in heterochromatin has been studied by in situ hybridisation to mitotic chromosomes.
The repressive effect of su(Hw) on y2 expression is limited to the chromosome in which the su(Hw) binding sites in gypsy are present. The negative effect of the su(Hw) protein can be transmitted to the gene present on the other homologous paired chromosome in the presence of mod(mdg4) mutations. They allow the su(Hw) protein to act in trans and inhibit the action of the y enhancers located in the homologous chromosome on the promoter of their gene.
Flies infected with the gypsy viral particles isolated from flam females show de novo gypsy mobilization accompanied by the appearance of new mutations. Reversion of various transposable
element-induced mutations was also observed at a high frequency. The presence of mutations correlates with the mobilization
of transposable elements such as copia, jockey, Stalker, 412 and Tirant.
The distribution of a number of transposable elements has been studied in 10 Harwich mutation accumulation lines.
The distribution of transposable elements within heterochromatin indicates that they are major structural components of the
heterochromatin.
22% of recessive lethal mutations caused by the insertion of P{SUPor-P}, which contains gypsy\su(Hw)BR sequences, are suppressed by mutations in su(Hw), indicating that they would not have been detected by a standard P-element insertion.
su(Hw) acts as a transcriptional activator of gypsy expression during development. Ecol\lacZ reporter gene constructs containing the su(Hw) binding region upstream suggest that su(Hw) activates the tissue-specific expression of gypsy at the level of transcription initiation. Analysis of specific su(Hw) mutant alleles on the expression of this reporter gene indicate that both the amino terminal acidic and the leucine zipper
domains of su(Hw) are essential for the proper regulation of gypsy expression in the larval tissues and adult ovaries. This data suggests that the su(Hw) protein interacts with other proteins through its acidic and leucine zipper domains to produce the tissue-specific expression
of gypsy.
The distribution of gypsy elements across the chromosomes has been analysed in individuals from a natural population of D.melanogaster.
The spatial and temporal expression patterns of fifteen families of retrotransposons are analysed during embryogenesis and
are found to be conserved. Results suggest that all families carry cis-acting elements that control their spatial and temporal
expression patterns.
gypsy can be transmitted by microinjection of egg plasm from embryos of a strain containing actively transposing gypsy elements into embryos of a strain originally devoid of transposing elements. gypsy is an infectious virus as shown by horizontal transfer when individuals devoid of transposing elements are raised on medium
containing ground pupae of a stock containing transposable elements.
Comparisons between gypsy sequences in D.melanogaster, D.subobscura and D.virilis strongly suggest that gypsy sequences have been horizontally transferred between these species.
Plasmids containing 5' truncated gypsy elements were introduced into D.melanogaster and D.hydei cell lines: appearance of new complete DNA copies with reconstructed 5'LTR were detected by PCR after transient expression
and by Southern blot after stable transformation. Two gypsy subfamilies that differ in transpositional activity were found to differ in efficiency of retrotransposition.
gypsy and Dvir\gypsy virus-like particles (VLP) are present in the cultured media from D.melanogaster and D.virilis.
Stability of 11 transposable element families compared by Southern blotting among individuals of lines that had been subjected
to 30 generations of sister sib matings. 412, roo, blood, 297, 1731 and G-element all appear stable, whereas copia, hobo, I-element, gypsy and jockey elements show instability.
Numbers of mdg1, mdg3, gypsy and copia have been studied in several strains of D.melanogaster and D.simulans. Mean number of mdg1 and copia sites are drastically reduced in D.simulans. Majority of mdg1 and copia sites, and one third of mdg3 sites, are in hot spots for insertion, particularly in D.simulans. Southern blot analysis indicates that the majority of mdg1 and copia are in the euchromatin of D.melanogaster but the heterochromatin of D.simulans.
gypsy contains a sequence, BaBx, that potentiates upstream polyadenylation sites and it appears to operate at the level of the
DNA template. Changes in the repeat motif can reduce both protein binding and poly(A) site potentiation properties of the
sequence.
The distribution of a number of transposable elements, including gypsy elements, in a D.melanogaster laboratory strain with a high frequency of spontaneous mutations and its derivatives, has been studied.
gypsy elements from two D.melanogaster strains (one stable and one unstable) have been cloned and analysed. The stable strain contains a small number of gypsy elements with a constant localisation in the chromosomes. The unstable strain contains an increased number of gypsy elements with a higher frequency of transposition.
The distribution of gypsy elements in a mutator strain of D.melanogaster and its derivatives has been studied.
gypsy elements from a single D.melanogaster strain are structurally heterogeneous, consisting of full-length, slightly diverging, and highly diverging gypsy elements.
The gypsy element contains two closely situated regions that bind to proteins from nuclear extracts. One is an imperfect palindrome
with homology to the E.coli lac operator and the other is a reiterated sequence with homology to the octamer sequence found in the core of many enhancers
and upstream promoter elements. Transient expression of deletion mutants shows that the sequences are negative and positive
regulators, respectively, of gypsy transcription. The su(Hw) gene encodes a protein which activates gypsy transcription and the su(f) gene encodes a protein capable of gypsy repression.
First described by Ilyin et al. (1980) as mdg4, a sequence complementary to double stranded RNA from tissue culture cells
and by Bender et al. (1983) as an insertion associated with mutations Ubxbx-3, Ubxbx-34e, Ubxbxd-55i and Ubxbxd-51j. Modolell et al. (1983) have shown by in situ hybridization that gypsy insertions are associated with many mutations suppressed
by su(Hw). The su(Hw) product binds to an enhancer-like sequence within gypsy and this may affect the expression of adjacent
genes as well as of gypsy itself. This is alleviated by su(Hw) mutations (Geyer et al., 1988; Peifer and Bender, 1988; Spana
et al., 1988; Mazo et al., 1989). The phenotypes of some mutations caused by gypsy insertions are affected by su(f) mutations.
The LTR sequence shown here was reported by Bayev et al. Freund and Meselson (1984) have reported an equivalent sequence of
482 base pairs. The sequences of complete elements have been reported by Yuki et al. and Marlor et al. (1986). The map shown
in Lindsley, Zimm, 1992: 1101 is based on those of Bayev et al. and Mattox and Davidson (1984). There are no sites for the
enzymes BamHI or SalI.
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Other Information
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Etymology
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External Crossreferences
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| Sequence Crossreferences | |||
| Other Crossreferences | |||
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Synonyms & Secondary IDs
( 17 )
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| Reported As | |||
| Symbol Synonym |
gypsy
(Heredia et al., 2004, Zamore, 2007, Mevel-Ninio et al., 2007, Saito et al., 2006, Abe et al., 2001, Zaratiegui, 2007, Vagin et al., 2006, Ramos et al., 2006, Akbari et al., 2006, Zhang et al., 2011, Ganko et al., 2006, Linheiro and Bergman, 2012, Nefedova and Kim, 2007, Semin et al., 2001, Pelisson et al., 2007, Salenko et al., 2007, Pane et al., 2011, Fablet et al., 2007, Minervini et al., 2007, Pai et al., 2004, Gunawardane et al., 2007, Lau et al., 2009, Gause et al., 2001, Gause and Georgiev, 2000, Petrov et al., 2011, Gabus et al., 2006, Brennecke et al., 2007, Savitsky et al., 2002, Georgiev et al., 2008, Walser et al., 2006, Mugnier et al., 2008, Steinbiss et al., 2009, Kotnova et al., 2009, Yu et al., 2011, Kulkarni and Arnosti, 2005, Kravchenko et al., 2005, Savitskaya et al., 2006, Golovnin et al., 2007, Liu et al., 2011, Nefedova et al., 2011, Matyunina et al., 2008, Brennecke et al., 2008, Czech et al., 2008, Deloger et al., 2009, Kawamura et al., 2008, Xu et al., 2004, Kuhn-Parnell et al., 2008, Labrador et al., 2008, Melnikova et al., 2008, Chetverina et al., 2008, Melnikova et al., 2008, Lerat et al., 2011, Anand and Kai, 2012, Malone et al., 2009, Tchurikov et al., 2009, Golovnin et al., 2008, Li et al., 2009, Lei and Corces, 2006, Ronfort et al., 2004, Malik and Henikoff, 2005, Fanti et al., 2003, Golovnin et al., 2008, Adryan et al., 2007, Patil and Kai, 2010, Kotova et al., 2010, Moshkovich and Lei, 2010, Oliver et al., 2010, Pelisson et al., 2007, Zamparini et al., 2011, Nefedova et al., 2006, Syomin and Ilyin, 2006, Tan et al., 2012, Sienski et al., 2012, Silicheva et al., 2010, Golovnin et al., 2012)
Gypsy
GYPSY_I
gypsy/mdg4
GYPSYMEL
MDG4
mdg4
(Bayev, 1994.2.28, Bayev, 1994.2.28, Bayev, 1994.2.28, Smirnova et al., 1998, Maksimiv et al., 1995, Golubovsky and Kaidanov, 1995, Georgiev and Kozycina, 1993, Kim and Belyaeva, 1991, Arkhipova and Ilyin, 1991, Leibovich, 1990, Georgiev and Gerasimova, 1989, Georgiev et al., 1989, Kim et al., 1989, FlyBase, 1996-, Nefedova et al., 2006)
mdg-4
MGE4
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| Name Synonym |
gypsy
gypsy element
gypsy retrotransposon
Gypsy virus
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| Secondary FlyBase IDs | |||
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References
( 363 )
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Recent research papers ( 16 )
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Recent reviews (0)
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| All reviews listed in FlyBase were published before 2011 | |||