1887

Chapter 21 : P Transposable Elements in Drosophila melanogaster

MyBook is a cheap paperback edition of the original book and will be sold at uniform, low price.

Preview this chapter:
Zoom in
Zoomout

P Transposable Elements in Drosophila melanogaster, Page 1 of 2

| /docserver/preview/fulltext/10.1128/9781555817954/9781555812096_Chap21-1.gif /docserver/preview/fulltext/10.1128/9781555817954/9781555812096_Chap21-2.gif

Abstract:

P transposable elements are one of the best studied eukaryotic mobile DNA elements in metazoans. This chapter focuses on the more recent developments in understanding the mechanism and specificity of P element transposition through detailed biochemical and genetic experiments. With the use of P element transformation to introduce the yeast site-specific Flip recombinase (FLP) and the Flip recombinase target (FRT) recombination sites into the genome, it was shown that the FLP recombinase could catalyze DNA strand exchange between FRT sites. Subsequently, it was shown that strand exchange could occur between FRT sites on different chromosomes or sister chromatids. With the development of efficient single P element mutagenesis strategies, many laboratories have undertaken large-scale P element insertional mutation screens. The ability to stimulate recombination at P element sites has had several useful applications in genetics. A system for purifying the P element transposase protein from tissue culture cell nuclear extracts was developed by using a combination of conventional and affinity chromotography methods. It will be extremely important and interesting for the future to understand how P element mobility might be linked to the cell cycle and DNA repair checkpoints and how P element transposition might be influenced by extracellular stimuli.

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21

Key Concept Ranking

Transcription Start Site
0.41725993
0.41725993
Highlighted Text: Show | Hide
Loading full text...

Full text loading...

Figures

Image of Figure 1
Figure 1

The genetics and symptoms of hybrid dysgenesis. The reciprocal crosses of hybrid dysgenesis are shown. Only when P strain males are mated toMstrain females is germ line development abnormal, because of high rates of P element transposition. Progeny from reciprocal M male by P female, P× P or M × M crosses are normal. M females give rise to eggs with a state permissive for P element transposition (M cytotype), whereas P females give rise to eggs with a state restrictive for P element transposition (P cytotype).

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 2
Figure 2

Features of the complete 2.9-kb P element. (A) Sequence features of the 2.9-kb P element. The four coding exons (ORF0, 1, 2, and 3) are indicated by boxes with nucleotide numbers shown. The positions of the three introns (IVS1, 2, 3) are indicated below. The DNA sequences of the terminal inverted 31-bp repeats and the internal 11-bp inverted repeats are shown, with corresponding nucleotide numbers shown above. The 8-bp duplications (dup) of target site DNA are shown by boxes at the ends of the element. (B) cis-acting elements of P element transposition. Key sequence features of the left (5′) and right (3′) end are indicated. The terminal 31-bp inverted repeats and the 11-bp internal inverted repeats are indicated by arrows. The transposase binding sites and 8-bp target site duplications are indicated by boxes. The distinct spacer lengths between the 31-bp repeats and the transposase binding sites, 21 bp at the 5′ end and 9 bp at the 3′ end, are indicated above.

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 3
Figure 3

P element mRNAs and proteins. The 2.9-kb P element and four exons (ORF0, 1, 2, and 3) are shown at the top. The germ line mRNA, in which all three introns are removed, encodes the 87-kDa transposase mRNA. The somatic mRNA, from which only the first two introns are removed (and which is also expressed in germ line and somatic cells), encodes the 66-kDa repressor mRNA. Shown at the bottom is a KP element, which contains an internal deletion. This truncated element encodes a 24-kDa repressor protein.

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 4
Figure 4

P element-mediated germ line transformation. Outline of the method for germ line transformation of Drosophila with P element vectors. Two plasmids, one encoding the P element transposase protein but lacking P element ends and the second plasmid carrying a foreign DNA segment and an eye color marker gene (wor ry) within P element ends, are injected into the posterior pole of preblastoderm embryos. The transposase plasmid enters nuclei of presumptive germ line cells, is expressed, and leads to transposition of the P element from the second plasmid into Drosophila germ line chromosomes. After development of the injected embryos (G generation), the surviving adults are mated to w or ry flies (G generation), and the progeny from this cross (G generation) are scored for restoration of wild-type eye color. The transformation frequency is typically ∼20% of the fertile G adults.

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 5
Figure 5

Features of the P element transposase protein. The N-terminal region contains a C2HC motif and basic region involved in site-specific DNA binding. Adjacent to this region are consensus sites for phosphorylation by the ATM family of DNA repair-checkpoint PI-related protein kinases. These sites, when mutated, affect transposase activity in vivo (Beall et al., unpublished). There are two dimerization regions adjacent to the N-terminal DNA binding domain: dimerization region I is a leucine zipper motif and dimerization region II is C-terminal to the leucine zipper but does not resemble any known motif. The central part of the protein contains a GTP binding region, with some sequence motifs found in the GTPase superfamily ( ). The C-terminal region is highly acidic and contains four acidic residues (D444, D528, E531, and D545) that, when mutated, block transposase activity in vivo and in vitro (Ahrens et al., unpublished).

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 6
Figure 6

Amino acid sequences of the P element transposase and 66-kDa repressor protein. Single letter code for the 751-aminoacid and 576-amino-acid transposase and 66-kDa proteins. The alternative C terminus of the 66-kDa protein (GMTNLKECVNKVIP) encoded by sequences from IVS is indicated below the corresponding transposase residues (amino acids 562 to 576). The C2HC DNA binding motif is indicated by boxed shadowed letters. The leucine zipper motif is indicated by boxed, outlined L residues. The three GTP binding region homologies are indicated by white letters on a black background. Potential phosphorylation sites for the ATM family of DNA repair-checkpoint PI3 protein kinases (S/TQ or QS/T) are indicated by white letters on a black background. Potential phosphorylation sites for cdc2 (TPHL and TPLQ) are indicated by highlighting and underlining. Acidic residues at the C terminus are underlined, and the four acidic residues that, when mutated, inactivate the protein for catalysis in vivo and in vitro are boxed and highlighted.

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 7
Figure 7

Similarities between P element transposase and GTPase superfamily members. Alignments of regions of P element transposase that bear some resemblance to known G proteins. The conserved motifs for phosphoryl binding and guanine specificity are indicated at the top. Amino acid numbers are given below for ras, T antigen, and P element transposase. The residue D379 that when changed to N switched the specificity from guanosine to xanthosine in P element transposase is indicated at the bottom. Reprinted from reference 139 with permission.

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 8
Figure 8

Binding sites for transposase on P element DNA. DNA sequences from the 5′ end (nt 48 to 68) and from the 3′ end (nt 2855 to 2871) that are protected from DNase I cleavage by P element transposase. Distance of the beginning of the 10-bp core sequence from the corresponding 31-bp terminal repeat is indicated at the right. A consensus site derived by comparing the two protected regions is shown at the bottom.

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 9
Figure 9

Pathway of DNA cleavage and joining during P element transposition. Shown at the top is the P element donor site with the target site duplications, 31-bp inverted repeats, and the 5′ and 3′ cleavage sites indicated. In the first step of transposition, donor cleavage, both ends of the P element are cleaved. The novel cleavage results in a 17-nt single-strand extension on the P element ends and leaves 17 nt from each P element inverted repeat attached to the donor DNA cleavage site. Once transposon excision occurs, the donor site can be repaired via an NHEJ pathway (shown to the right) or via an SDSA gap repair pathway (not shown) (142). The excised P element then selects a target site and, upon strand transfer, integrates into the donor site, generating a gapped intermediate, which upon repair completes integration, creating an 8-bp duplication of target DNA (bottom). Reprinted from reference 10 with permission.

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 11
Figure 11

DNA repair pathways used at the donor site after transposase-mediated P element excision. After cleavage of P element DNA by transposase, the Drosophila Ku subunits may associate with the single-strand extensions. These cleaved termini can undergo two fates. One involves NHEJ where the cleaved termini are brought together and after DNA polymerase, ligase, and possibly nuclease action rejoining occurs with the P element sequences and target site duplications present ( ). The second pathway involves gap repair via the SDSA pathway. Either with or without resection, these single-stranded 3′ extensions find homologous DNA sequences to use as templates for DNA repair synthesis. Copying can occur from either P element or flanking Drosophila genomic sequences as templates ( ).

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 10
Figure 10

Model for gap repair after P element excision. Homologous chromosomes or sister chromatids, which after undergoing P element excision leave a double-strand gap at the donor site. The homologous sequence then serves as a template for SDSA synthesis ( ). Completion of repair replaces the original P element with a newly synthesized copy. If this gap repair process were incomplete, internal deletions of the P element would result.

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 12
Figure 12

Molecular mechanisms for repression of P element transposition. Two effects of repressor proteins can affect repression: transcriptional (left) and posttranscriptional (right). (Left) The binding of repressor to a transposase site at the 5′ P element end blocks transcription from the P element promoter, reducing transposase mRNA synthesis. (Right) The binding of repressor to the transposase sites occludes transposase-DNA interactions required to initiate P element transposition.

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 13
Figure 13

Model for P element trans-silencing. Telomeric P element insertions in TAS repeats interact with heterochromatin proteins. After mating, the euchromatic lacZ reporter P element transgene can form a paired complexwith the P element at the telomere (bottom). This association results in transcriptional repression of the reporter, hence the term “trans-silencing.”

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 14
Figure 14

Model for P cytotype regulation. Maternal P cytotype provides several components to control P element activity. Maternal deposition of repressor protein or RNA leads to reduction of P element pre-mRNA synthesis, altering the ratio of repressor and transposase mRNAs, resulting in an autoregulatory loop ( ). The maternal genome contains some P elements that acquire a maternal imprint of heterochromatin proteins (clustered grey circles) that, in the zygote, can lead to trans-silencing of certain P element insertions. Production and binding of the repressor to P element DNA can inhibit transposase RNA synthesis (left) and block transposase binding to P element DNA (right).

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 15
Figure 15

Alternative splicing of the P element third intron (IVS3). (A) Diagram of two IVS3 splicing patterns observed in vivo. The 5′ exon (ORF2), 3′ exon (ORF3), and intron (IVS3) elements are shown schematically. The negative regulatory elements in the 5′ exon (F1 and F2) are indicated. The accurate 5′ (1947) and 3′ (2138) splice sites are shown. The minor alternative D2 5′ (2048) splice site and alternative 3′ splice site (2017) with the flanking microexon is indicated below ( ). (B) RNA sequence of the IVS3 region in the P element. The accurate 5′ and 3′ splice sites are indicated, as is the intron branchpoint (BP). Also underlined are the internal (D1, D2, D3, and D4) and upstream (F1 and F2) 5′ splice site-like (pseudo) sequences ( Table 2 ) ( ). (C) The 5′ exon negative regulatory element and mutations that activate IVS3 splicing. Sequence of the P element 5′ exon. The F1 and F2 sites (labeled brackets) and the accurate 5′ splice site (arrowhead) are shown. Below is shown a reiterated sequence (PyAGNUUAAG) present overlapping the F1 and F2 sites. Below are shown 5′ exon mutations that activate IVS3 splicing in vitro ( ) or in vivo ( ).

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 16
Figure 16

Model for somatic inhibition of IVS splicing. U1 snRNP usually interacts with the IVS3 5′ splice site during the early steps of intron recognition and spliceosome assembly. In somatic cells (and in vitro) this site is blocked ( ). Mutations in the upstream negative regulatory element lead to activation of IVS3 splicing in vivo ( ) and in vitro ( ). The F1 site binds U1 snRNP ( ) and the F2 site binds the hnRNP protein, hrp48 ( ). An RNA-binding protein containing four KH domains that is expressed highly in somatic cells, called PSI, has also been implicated in IVS3 splicing control ( ). Several other uncharacterized proteins from Drosophila extracts have been identified by UV cross-linking to the IVS3 5′ exon ( ).

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Permissions and Reprints Request Permissions
Download as Powerpoint

References

/content/book/10.1128/9781555817954.chap21
1. Adams, M. D.,, S. E. Celniker,, R. A. Holt,, C. A. Evans,, J. D. Gocayne,, P. G. Amanatides,, S. E. Scherer,, P. W. Li,, R. A. Hoskins,, R. F. Galle,, R. A. George,, S. E. Lewis,, S. Richards,, M. Ashburner,, S. N. Henderson,, G. G. Sutton,, J. R. Wortman,, M. D. Yandell,, Q. Zhang,, L. X. Chen,, R. C. Brandon,, Y. H. Rogers,, R. G. Blazej,, M. Champe,, B. D. Pfeiffer,, K. H. Wan,, C. Doyle,, E. G. Baxter,, G. Helt,, C. R. Nelson,, G. L. Gabor Miklos,, J. F. Abril,, A. Agbayani,, H. J. An,, C. Andrews- Pfannkoch,, D. Baldwin,, R. M. Ballew,, A. Basu,, J. Baxendale,, L. Bayraktaroglu,, E. M. Beasley,, K. Y. Beeson,, P. V. Benos,, B. P. Berman,, D. Bhandari,, S. Bolshakov,, D. Borkova,, M. R. Botchan,, J. Bouck,, P. Brokstein,, P. Brottier,, K. C. Burtis,, D. A. Busam,, H. Butler,, E. Cadieu,, A. Center,, I. Chandra,, J. M. Cherry,, S. Cawley,, C. Dahlke,, L. B. Davenport,, P. Davies,, B. de Pablos,, A. Delcher,, Z. Deng,, A. D. Mays,, I. Dew,, S. M. Dietz,, K. Dodson,, L. E. Doup,, M. Downes,, S. Dugan- Rocha,, B. C. Dunkov,, P. Dunn,, K. J. Durbin,, C. C. Evangelista,, C. Ferraz,, S. Ferriera,, W. Fleischmann,, C. Fosler,, A. E. Gabrielian,, N. S. Garg,, W. M. Gelbart,, K. Glasser,, A. Glodek,, F. Gong,, J. H. Gorrell,, Z. Gu,, P. Guan,, M. Harris,, N. L. Harris,, D. Harvey,, T. J. Heiman,, J. R. Hernandez,, J. Houck,, D. Hostin,, K. A. Houston,, T. J. Howland,, M. H. Wei,, C. Ibegwam,, M. Jalali,, F. Kalush,, G. H. Karpen,, Z. Ke,, J. A. Kennison,, K. A. Ketchum,, B. E. Kimmel,, C. D. Kodira,, C. Kraft,, S. Kravitz,, D. Kulp,, Z. Lai,, P. Lasko,, Y. Lei,, A. A. Levitsky,, J. Li,, Z. Li,, Y. Liang,, X. Lin,, X. Liu,, B. Mattei,, T. C. McIntosh,, M. P. McLeod,, D. McPherson,, G. Merkulov,, N. V. Milshina,, C. Mobarry,, J. Morris,, A. Moshrefi,, S. M. Mount,, M. Moy,, B. Murphy,, L. Murphy,, D. M. Muzny,, D. L. Nelson,, D. R. Nelson,, K. A. Nelson,, K. Nixon,, D. R. Nusskern,, J. M. Pacleb,, M. Palazzolo,, G. S. Pittman,, S. Pan,, J. Pollard,, V. Puri,, M. G. Reese,, K. Reinert,, K. Remington,, R. D. Saunders,, F. Scheeler,, H. Shen,, B. C. Shue,, I. Siden- Kiamos,, M. Simpson,, M. P. Skupski,, T. Smith,, E. Spier,, A. C. Spradling,, M. Stapleton,, R. Strong,, E. Sun,, R. Svirskas,, C. Tector,, R. Turner,, E. Venter,, A. H. Wang,, X. Wang,, Z. Y. Wang,, D. A. Wassarman,, G. M. Weinstock,, J. Weissenbach,, S. M. Williams,, T. Woodage,, K. C. Worley,, D. Wu,, S. Yang,, Q. A. Yao,, J. Ye,, R. F. Yeh,, J. S. Zaveri,, M. Zhan,, G. Zhang,, Q. Zhao,, L. Zheng,, X. H. Zheng,, F. N. Zhong,, W. Zhong,, X. Zhou,, S. Zhu,, X. Zhu,, H. O. Smith,, R. A. Gibbs,, E. W. Myers,, G. M. Rubin,, and J. C. Venter. 2000. The genome sequence of Drosophila melanogaster. Science 287: 21852195.
2. Adams, M. D.,, R. S. Tarng,, and D. C. Rio. 1997. The alternative splicing factor PSI regulates P-element third intron splicing in vivo. Genes Dev. 11:129138.
3. Andrews, J. D.,, and G. B. Gloor. 1995. A role for the KP leucine zipper in regulating P element transposition in Drosophila melanogaster. Genetics 141:587594.
4. Ashburner, M.,, S. Misra,, J. Roote,, S. E. Lewis,, R. Blazej,, T. Davis,, C. Doyle,, R. Galle,, and N. Harris. 1999. An exploration of the sequence of a 2.9-Mb region of the genome of Drosophila melanogaster. Genetics 153:179219.
5. Ballinger, D. G.,, and S. Benzer. 1989. Targeted gene mutations in Drosophila. Proc. Natl. Acad. Sci. USA 86: 94029406.
6. Banga, S. S.,, A. Velazquez,, and J. B. Boyd. 1991. P transposition in Drosophila provides a new tool for analyzing postreplication repair and double-strand break repair. Mutat. Res. 255:7988.
7. Basler, K.,, and G. Struhl. 1994. Compartment boundaries and the control of Drosophila limb pattern by hedgehog protein. Nature 368:208214.
8. Beall, E. L.,, A. Admon,, and D. C. Rio. 1994. A Drosophila protein homologous to the human p70 Ku autoimmune antigen interacts with the P transposable element inverted repeats. Proc. Natl. Acad. Sci. USA 91:1268112685.
9. Beall, E. L.,, and D. C. Rio. 1996. Drosophila IRBP/Ku p70 corresponds to the mutagen-sensitive mus309 gene and is involved in P-element excision in vivo. Genes Dev. 10: 921933.
10. Beall, E. L.,, and D. C. Rio. 1997. Drosophila P-element transposase is a novel site-specific endonuclease. Genes Dev. 11: 21372151.
11. Beall, E. L.,, and D. C. Rio. 1998. Transposase makes critical contacts with, and is stimulated by, single-stranded DNA at the P element termini in vitro. EMBO J. 17:21222136.
12. Bellaiche, Y.,, V. Mogila,, and N. Perrimon. 1999. I-Scel endonuclease, a new tool for studying DNA double-strand break repair mechanisms in Drosophila. Genetics 152:10371044.
13. Bellen, H. J.,, C. J. O’Kane,, C. Wilson,, U. Grossniklaus,, R. K. Pearson,, and W. J. Gehring. 1989. P-element-mediated enhancer detection: a vesatile method to study development in Drosophila. Genes Dev. 3:12881300.
14. Berg, C. A.,, and A. C. Spradling. 1991. Studies on the rate and site-specificity of P element transposition. Genetics 127: 515524.
15. Bier, E.,, H. Vaessin,, S. Shepherd,, K. Lee,, K. McCall,, S. Barbel,, L. Ackerman,, R. Carretto,, T. Uemura,, E. Grell, et al. 1989. Searching for pattern and mutation in the Drosophila genome with a P-lacZ vector. Genes Dev. 3:12731287.
16. Bingham, P. M.,, M. G. Kidwell,, and G. M. Rubin. 1982. The molecular basis of P-M hybrid dysgenesis: the role of the P element, a P strain-specific transposon family. Cell 29: 9951004.
17. Bingham, P. M.,, R. Levis,, and G. M. Rubin. 1981. Cloning of DNA sequences from the white locus of D. melanogaster by a novel and general method. Cell 25:693704.
18. Black, D. M.,, M. S. Jackson,, M. G. Kidwell,, and G. A. Dover. 1987. KP elements repress P-induced hybrid dysgenesis in Drosophila melanogaster. EMBO J. 6:41254135.
19. Bourne, H. R.,, D. A. Sanders,, and F. McCormick. 1990. The GTPase superfamily: a conserved switch for diverse cell functions. Nature 348:125132.
20. Bourne, H. R.,, D. A. Sanders,, and F. McCormick. 1991. The GTPase superfamily: conserved structure and molecular mechanism. Nature 349:117127.
21. Boyd, J. B.,, M. D. Golino,, K. E. Shaw,, C. J. Osgood,, and M. M. Green. 1981. Third-chromosome mutagen-sensitive mutants of Drosophila melanogaster. Genetics 97:607623.
22. Boyd, J. B.,, and R. B. Setlow. 1976. Characterization of postreplication repair in mutagen-sensitive strains of Drosophila melanogaster. Genetics 84:507526.
23. Brand, A. H.,, A. S. Manoukian,, and N. Perrimon. 1994. Ectopic expression in Drosophila. Methods Cell Biol. 44: 635654.
24. Brand, A. H.,, and N. Perrimon. 1993. Targeted gene expression as a means of altering cell fates and generating dominant phenotypes. Development 118:401415.
25. Brautigam, C. A.,, and T. A. Steitz. 1998. Structural and functional insights provided by crystal structures of DNA polymerases and their substrate complexes. Curr. Opin. Struct. Biol. 8:5463.
26. Burd, C. G.,, and G. Dreyfuss. 1994. RNA binding specificity of hnRNP A1: significance of hnRNP A1 high-affinity binding sites in pre-mRNA splicing. EMBO J. 13:11971204.
27. Burd, C. G.,, and G. Dreyfuss. 1994. Conserved structures and diversity of functions of RNA-binding proteins. Science 265:615621.
28. Cech, T. R. 1990. Self-splicing of group I introns. Annu. Rev. Biochem. 59:543568.
29. Chabot, B. 1996. Directing alternative splicing: cast and scenarios. Trends Genet. 12:472478.
30. Chain, A. C.,, S. Zollman,, J. C. Tseng,, and F. A. Laski. 1991. Identification of a cis-acting sequence required for germ linespecific splicing of the P element ORF2-ORF3 intron. Mol. Cell. Biol. 11:15381546.
31. Chen, B.,, T. Chu,, E. Harms,, J. P. Gergen,, and S. Strickland. 1998. Mapping of Drosophila mutations using site-specific male recombination. Genetics 149:157163.
32. Chou, T. B.,, E. Noll,, and N. Perrimon. 1993. Autosomal P[ovoD1] dominant female-sterile insertions in Drosophila and their use in generating germ-line chimeras. Development 119:13591369.
33. Chou, T. B.,, and N. Perrimon. 1992. Use of a yeast sitespecific recombinase to produce female germline chimeras in Drosophila. Genetics 131:643653.
34. Chou, T. B.,, and N. Perrimon. 1996. The autosomal FLPDFS technique for generating germline mosaics in Drosophila melanogaster. Genetics 144:16731679.
35. Chow, S. A.,, K. A. Vincent,, V. Ellison,, and P. O. Brown. 1992. Reversal of integration and DNA splicing mediated by integrase of human immunodeficiency virus. Science 255: 723726.
36. Chu, G. 1997. Double strand break repair. J. Biol. Chem. 272:2409724100.
37. Coen, D. 1990. P element regulatory products enhance zeste repression of a P[white duplicated] transgene in Drosophila melanogaster. Genetics 126:949960.
38. Cooley, L.,, R. Kelley,, and A. Spradling. 1988. Insertional mutagenesis of the Drosophila genome with single P elements. Science 239:11211128.
39. Cooley, L.,, D. Thompson,, and A. C. Spradling. 1990. Constructing deletions with defined endpoints in Drosophila. Proc. Natl. Acad. Sci. USA 87:31703173.
40. Craig, N. L. 1995. Unity in transposition reactions. Science 270:253254.
41. Critchlow, S. E.,, and S. P. Jackson. 1998. DNA end-joining: from yeast to man. Trends Biochem. Sci. 23:394398.
42. Curcio, M. J.,, and D. J. Garfinkel. 1999. New lines of host defense: inhibition of Ty1 retrotransposition by Fus3p and NER/TFIIH. Trends Genet. 15:4345.
43. Dalby, B.,, A. J. Pereira,, and L. S. Goldstein. 1995. An inverse PCR screen for the detection of P element insertions in cloned genomic intervals in Drosophila melanogaster. Genetics 139: 757766.
44. Dang, D. T.,, and N. Perrimon. 1992. Use of a yeast sitespecific recombinase to generate embryonic mosaics in Drosophila. Dev. Genet 13:367375.
45. Desai, A.,, and T. J. Mitchison. 1997. Microtubule polymerization dynamics. Annu. Rev. Cell. Dev. Biol. 13:83117.
46. Dray, T.,, and G. B. Gloor. 1997. Homology requirements for targeting heterologous sequences during P-induced gap repair in Drosophila melanogaster. Genetics 147:689699.
47. Dynan, W. S.,, and S. Yoo. 1998. Interaction of Ku protein and DNA-dependent protein kinase catalytic subunit with nucleic acids. Nucleic Acids Res. 26:15511559.
48. Engels, W. R. 1979. Hybrid dysgenesis in Drosophila melanogaster: rules of inheritance of female sterility. Genet. Res. 33: 219236.
49. Engels, W. R. 1979. Extrachromosomal control of mutability in Drosophila melanogaster. Proc. Natl. Acad. Sci. USA 76: 40114015.
50. Engels, W. R. 1983. The P family of transposable elements in Drosophila. Annu. Rev. Genet. 17:315344.
51. Engels, W. R. 1984. A trans-acting product needed for P factor transposition in Drosophila. Science 226:11941196.
52. Engels, W. R., 1989. P elements in Drosophila melanogaster, p. 437484. In D. E. Berg, and M. M. Howe (ed.), Mobile DNA. American Society for Microbiology, Washington, D.C..
53. Engels, W. R. 1992. The origin of P elements in Drosophila melanogaster. Bioessays 14:681686.
54. Engels, W. R. 1996. P elements in Drosophila. Curr. Top. Microbiol. Immunol. 204:103123.
55. Engels, W. R. 1997. Invasions of P elements. Genetics 145: 1115.
56. Engels, W. R.,, W. K. Benz,, C. R. Preston,, P. L. Graham,, R. W. Phillis,, and H. M. Robertson. 1987. Somatic effects of P element activity in Drosophila melanogaster: pupal lethality. Genetics 117:745757.
57. Engels, W. R.,, D. M. Johnson-Schlitz,, W. B. Eggleston,, and J. Sved. 1990. High-frequency P element loss in Drosophila is homolog-dependent. Cell 62:515525.
58. Engels, W. R.,, C. R. Preston,, and D. M. Johnson-Schlitz. 1994. Long-range cis preference in DNA homology search over the length of a Drosophila chromosome. Science 263: 16231625.
59. Esposito, D.,, and R. Craigie. 1999. HIV integrase structure and function. Adv. Virus Res. 52:319333.
60. Fire, A. 1999. RNA-triggered gene silencing. Trends Genet. 15:358363.
61. Formosa, T.,, and B. M. Alberts. 1986. DNA synthesis dependent on genetic recombination: characterization of a reaction catalyzed by purified bacteriophage T4 proteins. Cell 47: 793806.
62. Fugmann, S. D.,, I. J. Villey,, L. M. Ptaszek,, and D. G. Schatz. 2000. Identification of two catalytic residues in RAG1 that define a single active site within the RAG1/RAG2 protein complex. Mol. Cell 5:97107.
63. Gell, D.,, and S. P. Jackson. 1999. Mapping of protein-protein interactions within the DNA-dependent protein kinase complex. Nucleic Acids Res. 27:34943502.
64. Gloor, G. B.,, and D. H. Lankenau. 1998. Gene conversion in mitotically dividing cells: a view from Drosophila. Trends Genet. 14:4346.
65. Gloor, G. B.,, J. Moretti,, J. Mouyal,, and K. J. Keeler. 2000. Distinct P element excision products in somatic and germline cells of Drosophila. Genetics 155:18211830.
66. Gloor, G. B.,, N. A. Nassif,, D. M. Johnson-Schlitz,, C. R. Preston,, and W. R. Engels. 1991. Targeted gene replacement in Drosophila via P element-induced gap repair. Science 253: 11101117.
67. Gloor, G. B.,, C. R. Preston,, D. M. Johnson-Schlitz,, N. A. Nassif,, R. W. Phillis,, W. K. Benz,, H. M. Robertson,, and W. R. Engels. 1993. Type I repressors of P element mobility. Genetics 135:8195.
68. Golic, K. G. 1991. Site-specific recombination between homologous chromosomes in Drosophila. Science 252: 958961.
69. Golic, K. G.,, and M. M. Golic. 1996. Engineering the Drosophila genome: chromosome rearrangements by design. Genetics 144:16931711.
70. Golic, K. G.,, and S. Lindquist. 1989. The FLP recombinase of yeast catalyzes site-specific recombination in the Drosophila genome. Cell 59:499509.
71. Gray, Y. H. M.,, M. M. Tanaka,, and J. A. Sved. 1996. P element-induced recombination in Drosophila melanogaster: hybrid element insertion. Genetics 144:16011610.
72. Grindley, N. D.,, and A. E. Leschziner. 1995. DNA transposition: from a black boxto a color monitor. Cell 83: 10631066.
73. Grossniklaus, U.,, H. J. Bellen,, C. Wilson,, and W. J. Gehring. 1989. P-element-mediated enhancer detection applied to the study of oogenesis in Drosophila. Genes Dev. 107:189200.
74. Hamilton, B. A.,, M. J. Palazzolo,, J. H. Chang,, K. VijayRaghavan,, C. A. Mayeda,, M. A. Whitney,, and E. M. Meyerowitz. 1991. Large scale screen for transposon insertions into cloned genes. Proc. Natl. Acad. Sci. USA 88:27312735.
75. Hamilton, B. A.,, and K. Zinn. 1994. From clone to mutant gene. Methods Cell Biol. 44:8194.
76. Hammond, L. E.,, D. Z. Rudner,, R. Kanaar,, and D. C. Rio. 1997. Mutations in the hrp48 gene, which encodes a Drosophila heterogeneous nuclear ribonucleoprotein particle protein, cause lethality and developmental defects and affect Pelement third-intron splicing in vivo. Mol. Cell Biol. 17: 72607267.
77. Hay, B. A.,, R. Maile,, and G. M. Rubin. 1997. P element insertion-dependent gene activation in the Drosophila eye. Proc. Natl. Acad. Sci. USA 94:51955200.
78. Hazelrigg, T.,, R. Levis,, and G. M. Rubin. 1984. Transforma tion of white locusDNAin drosophila: dosage compensation, zeste interaction, and position effects. Cell 36:469481.
79. Heath, E. M.,, and M. J. Simmons. 1991. Genetic and molecular analysis of repression in the P-M system of hybrid dysgenesis in Drosophila melanogaster. Genet. Res. 57:213226.
80. Hesse, J. E.,, M. R. Lieber,, M. Gellert,, and K. Mizuuchi. 1987. Extrachromosomal DNA substrates in pre-B cells undergo inversion or deletion at immunoglobulin V-(D)-J joining signals. Cell 49:775783.
81. Hickman, A. B.,, Y. Li,, S. V. Mathew,, E. W. May,, N. L. Craig,, and F. Dyda. 2000. Unexpected structural diversity in DNA recombination: the restriction endonuclease connection. Mol. Cell 5:1002511034.
82. Hiraizumi, Y. 1971. Spontaneous recombination in Drosophila melanogaster males. Proc. Natl. Acad. Sci. USA 68: 268270.
83. Hirose, Y.,, and J. L. Manley. 2000. RNA polymerase II and the integration of nuclear events. Genes Dev. 14:14151429.
84. Horowitz, D. S.,, and A. R. Krainer. 1994. Mechanisms for selecting 5′ splice sites in mammalian pre-mRNA splicing. Trends Genet. 10:100106.
85. Hoskins, R. A.,, C. R. Nelson,, B. P. Berman,, T. R. Laverty,, R. A. George,, L. Ciesiolka,, M. Naeemuddin,, A. D. Arenson,, J. Durbin,, R. G. David,, P. E. Tabor,, M. R. Bailey,, D. R. DeShazo,, J. Catanese,, A. Mammoser,, K. Osoegawa,, P. J. de Jong,, S. E. Celniker,, R. A. Gibbs,, G. M. Rubin,, and S. E. Scherer. 2000. A BAC-based physical map of the major autosomes of Drosophila melanogaster. Science 287:22712274.
86. Houck, M. A.,, J. B. Clark,, K. R. Peterson,, and M. G. Kidwell. 1991. Possible horizontal transfer of Drosophila genes by the mite Proctolaelaps regalis. Science 253:11251128.
87. Jackson, M. S.,, D. M. Black,, and G. A. Dover. 1988. Amplification of KP elements associated with the repression of hybrid dysgenesis in Drosophila melanogaster. Genetics 120: 10031013.
88. Jacoby, D. B.,, and P. C. Wensink. 1994. Yolk protein factor 1 is a Drosophila homolog of Ku, the DNA-binding subunit of a DNA-dependent protein kinase from humans. J. Biol. Chem. 269:1148411491.
89. Jasin, M. 1996. Genetic manipulation of genomes with rarecutting endonucleases. Trends Genet. 12:224228.
90. Johnson-Schlitz, D. M.,, and W. R. Engels. 1993. P-elementinduced interallelic gene conversion of insertions and deletions in Drosophila melanogaster. Mol. Cell. Biol. 13: 70067018.
91. Joyce, C. M.,, and T. A. Steitz. 1995. Polymerase structures and function: variations on a theme? J. Bacteriol. 177: 63216329.
92. Kaiser, K.,, and S. F. Goodwin. 1990. ‘‘Site-selected’’ transposon mutagenesis of Drosophila. Proc. Natl. Acad. Sci. USA 87:16861690.
93. Kanaar, R.,, J. H. Hoeijmakers,, and D. C. van Gent. 1998. Molecular mechanisms of DNA double strand break repair. Trends Cell. Biol. 8:483489.
94. Karess, R. E.,, and G. M. Rubin. 1984. Analysis of P transposable element function in Drosophila. Cell 38:135146.
95. Karpen, G. H.,, and A. C. Spradling. 1992. Analysis of subtelomeric heterochromatin in the Drosophila minichromosome Dp1187 by single P element insertional mutagenesis. Genetics 132:737753.
96. Kassis, J. A.,, E. Noll,, E. P. VanSickle,, W. F. Odenwald,, and N. Perrimon. 1992. Altering the insertional specificity of a Drosophila transposable element. Proc. Natl. Acad. Sci. USA 89:19191923.
97. Kaufman, P. D.,, R. F. Doll,, and D. C. Rio. 1989. Drosophila P element transposase requires internal P element DNA sequences. Cell 38:135146.
98. Kaufman, P. D.,, and D. C. Rio. 1991. Drosophila P-element transposase is a transcriptional repressor in vitro. Proc. Natl. Acad. Sci. USA 88:26132617.
99. Kaufman, P. D.,, and D. C. Rio. 1992. P element transposition in vitro proceeds by a cut-and-paste mechanism and uses GTP as a cofactor. Cell 69:2739.
100. Keeler, K. J.,, T. Dray,, J. E. Penney,, and G. B. Gloor. 1996. Gene targeting of a plasmid-borne sequence to a doublestrand DNA break in Drosophila melanogaster. Mol. Cell. Biol. 16:522528.
101. Keeler, K. J.,, and G. B. Gloor. 1997. Efficient gap repair in Drosophila melanogaster requires a maximum of 31 nucleotides of homologous sequence at the searching ends. Mol. Cell. Biol. 17:627634.
102. Ketting, R. F.,, T. H. Haverkamp,, H. G. van Luenen,, and R. H. Plasterk. 1999. Mut-7 of C. elegans, required for transposon silencing and RNA interference, is a homolog of Werner syndrome helicase and RNaseD. Cell 99:133141.
103. Kidwell, M. G. 1981. Hybrid dysgenesis in Drosphila melanogaster: the genetics of cytotype determination in a neutral strain. Genetics 98:275290.
104. Kidwell, M. G. 1983. Evolution of hybrid dysgenesis determinants in Drosophila melanogaster. Proc. Natl. Acad. Sci. USA 80:16551659.
105. Kidwell, M. G. 1985. Hybrid dysgenesis in Drosophila melanogaster: nature and inheritance of P element regulation. Genetics 111:337350.
106. Kidwell, M. G. 1992. Horizontal transfer of P elements and other short inverted repeat transposons. Genetica 86: 275286.
107. Kidwell, M. G. 1992. Horizontal transfer. Curr. Opin. Genet. Dev. 2:868873.
108. Kidwell, M. G.,, and J. F. Kidwell. 1975. Cytoplasm-chromosome interactions in Drosophila melanogaster. Nature 253: 755756.
109. Kidwell, M. G.,, J. F. Kidwell,, and M. Nei. 1973. A case of high rate of spontaneous mutation affecting viability in Drosophila melanogaster. Genetics 75:133153.
110. Kidwell, M. G.,, J. F. Kidwell,, and J. A. Sved. 1977. Hybrid dysgenesis in Drosophila melanogaster: a syndrome of aberrant traits including mutation, sterility and male recombination. Genetics 86:813833.
111. Kidwell, M. G.,, and J. B. Novy. 1979. Hybrid dysgenesis in Drosophila melanogaster: sterility resulting from gonadal dysgenesis in the P-M system. Genetics 92:11271140.
112. Kim, D. R.,, Y. Dai,, C. L. Mundy,, W. Yang,, and M. A. Oettinger. 1999. Mutations of acidic residues in RAG1 define the active site of the V(D)J recombinase. Genes Dev. 13: 30703080.
113. Kim, S. T.,, D. S. Lim,, C. E. Canman,, and M. B. Kastan. 1999. Substrate specifications and identification of putative substrates of ATM kinase family members. J. Biol. Chem. 274:3753837543.
114. Kooistra, R.,, A. Pastink,, J. B. Zonneveld,, P. H. Lohman,, and J. C. Eeken. 1999. The Drosophila melanogaster DmRAD54 gene plays a crucial role in double- strand break repair after P-element excision and acts synergistically with Ku70 in the repair of X-ray damage. Mol. Cell. Biol. 19:62696275.
115. Krasnow, M. A.,, S. Cumberledge,, G. Manning,, L. A. Herzenberg,, and G. P. Nolan. 1991. Whole animal cell sorting of Drosophila embryos. Science 251:8185.
116. Kusano, K.,, D. M. Johnson-Schiltz,, and W. R. Engels. 2001. Sterility of Drosophila with mutations in the Bloom syndrome gene-complementation by Ku70. Science 291:26002602.
117. Labourier, E.,, M. D. Adams,, and D. C. Rio. 2001. Modulation of P element pre-mRNA splicing by a direct interaction between PSI and U1 snRNP 70K protein. Mol. Cell 8: 363373.
118. Landree, M. A.,, J. A. Wibbenmeyer,, and D. B. Roth. 1999. Mutational analysis of RAG1 and RAG2 identifies three catalytic amino acids in RAG1 critical for both cleavage steps of V(D)J recombination. Genes Dev. 13:30593069.
119. Lankenau, D. H.,, V. G. Corces,, and W. R. Engels. 1996. Comparison of targeted-gene replacement frequencies in Drosophila melanogaster at the forked and white loci. Mol. Cell. Biol. 16:35353544.
120. Lankenau, D. H.,, and G. B. Gloor. 1998. In vivo gap repair in Drosophila: a one-way street with many destinations. Bioessays 20:317327.
121. Laski, F. A.,, D. C. Rio,, and G. M. Rubin. 1986. Tissue specificity of Drosophila P element transposition is regulated at the level of mRNA splicing. Cell 44:719.
122. Laski, F. A.,, and G. M. Rubin. 1989. Analysis of the cisacting requirements for germ-line-specific splicing of the Pelement ORF2-ORF3 intron. Genes Dev. 3:720728.
123. Lee, C. C.,, E. L. Beall,, and D. C. Rio. 1998. DNA binding by the KP repressor protein inhibits P element transposase activity in vitro. EMBO J. 17:41664174.
124. Lee, C. C.,, Y. M. Mul,, and D. C. Rio. 1996. The Drosophila P-element KP repressor protein dimerizes and interacts with multiple sites on P-element DNA. Mol. Cell. Biol. 16: 56165622.
125. Lemaitre, B.,, and D. Coen. 1991. P regulatory products repress in vivo the P promoter activity in P-lacZ fusion genes. Proc. Natl. Acad. Sci. USA 88:44194423.
126. Lemaitre, B.,, S. Ronsseray,, and D. Coen. 1993. Maternal repression of the P element promoter in the germline of Drosophila melanogaster: a model for the P cytotype. Genetics 135:149160.
127. Liao, G. C.,, E. J. Rehm,, and G. M. Rubin. 2000. Insertion site preferences of the P transposable element in Drosophila melanogaster. Proc. Natl. Acad. Sci. USA 97:33473351.
128. Lopez, A. J. 1998. Alternative splicing of pre-mRNA: developmental consequences and mechanisms of regulation. Annu. Rev. Genet. 32:279305.
129. Matunis, E. L.,, M. J. Matunis,, and G. Dreyfuss. 1992. Characterization of the major hnRNP proteins from Drosophila melanogaster. J. Cell Biol. 116:257269.
130. Mayeda, A.,, and A. R. Krainer. 1992. Regulation of alternative pre-mRNA splicing by hnRNP A1 and splicing factor SF2. Cell 68:365375.
131. McClintock, B. 1984. The significance of responses of the genome to challenge. Science 226:792801.
132. Melek, M.,, and M. Gellert. 2000. RAG1/2-mediated resolution of transposition intermediates: two pathways and possible consequences. Cell 101:625633.
133. Miklos, G. L.,, and G. M. Rubin. 1996. The role of the genome project in determining gene function: insights from model organisms. Cell 86:521529.
134. Miller, W. J.,, N. Paricio,, S. Hagemann,, M. J. Martinez-Sebastian,, W. Pinsker,, and R. de Frutos. 1995. Structure and expression of clustered P element homologues in Drosophila subobscura and Drosophila guanche. Gene 156:167174.
135. Min, H.,, C. W. Turck,, J. M. Nikolic,, and D. L. Black. 1997. A new regulatory protein, KSRP, mediates exon inclusion through an intronic splicing enhancer. Genes Dev. 11: 10231036.
136. Misra, S.,, R. M. Buratowski,, T. Ohkawa,, and D. C. Rio. 1993. Cytotype control of Drosophila melanogaster P element transposition: genomic position determines maternal repression. Genetics 135:785800.
137. Misra, S.,, and D. C. Rio. 1990. Cytotype control of Drosophila P element transposition: the 66 kd protein is a repressor of transposase activity. Cell 62:269284.
138. Moore, M. J.,, C. C. Query,, and P. A. Sharp. 1993. Splicing of precursors to messenger RNAs by the spliceosome. In RNA World. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, N.Y..
139. Mul, Y. M.,, and D. C. Rio. 1997. Reprogramming the purine nucleotide cofactor requirement of Drosophila P element transposase in vivo. EMBO J. 16:44414447.
140. Mullins, M. C.,, D. C. Rio,, and G. M. Rubin. 1989. Cis-acting DNA sequence requirements for P-element transposition. Genes Dev. 3:729738.
141. Myers, E. W.,, G. G. Sutton,, A. L. Delcher,, I. M. Dew,, D. P. Fasulo,, M. J. Flanigan,, S. A. Kravitz,, C. M. Mobarry,, K. H. Reinert,, K. A. Remington,, E. L. Anson,, R. A. Bolanos,, H. H. Chou,, C. M. Jordan,, A. L. Halpern,, S. Lonardi,, E. M. Beasley,, R. C. Brandon,, L. Chen,, P. J. Dunn,, Z. Lai,, Y. Liang,, D. R. Nusskern,, M. Zhan,, Q. Zhang,, X. Zheng,, G. M. Rubin,, M. D. Adams,, and J. C. Venter. 2000. A whole-genome assembly of Drosophila. Science 287:21962204.
142. Nassif, N.,, J. Penney,, S. Pal,, W. R. Engels,, and G. B. Gloor. 1994. Efficient copying of nonhomologous sequences from ectopic sites via P-element-induced gap repair. Mol. Cell. Biol. 14:16131625.
143. Nitasaka, E.,, T. Mukai,, and T. Yamazaki. 1987. Repressor of P elements in Drosophila melanogaster: Cytotype determination by a defective P element carrying only open reading frames 0 through 2. Proc. Natl. Acad. Sci. USA 84: 76057608.
144. O’Brochta, D. A.,, S. P. Gomez,, and A. M. Handler. 1991. P element excision in Drosophila melanogaster and related drosophilids. Mol. Gen. Genet. 225:387394.
145. O’Hare, K.,, A. Driver,, S. McGrath,, and D. M. Johnson- Schiltz. 1992. Distribution and structure of cloned P elements from the Drosophila melanogaster P strain pi 2. Genet. Res. 60:3341.
146. O’Hare, K.,, and G. M. Rubin. 1983. Structures of P transposable elements and their sites of insertion and excision in the Drosophila melanogaster genome. Cell 34:2535.
147. O’Kane, C. J.,, and W. J. Gehring. 1987. Detection in situ of genomic regulatory elements in Drosophila. Proc. Natl. Acad. Sci. USA 84:91239127.
148. Paques, F.,, and J. E. Haber. 1999. Multiple pathways of recombination induced by double-strand breaks in Saccharomyces cerevisiae. Microbiol. Mol. Biol. Rev. 63:349404.
149. Paterson, J.,, and K. O’Hare. 1991. Structure and transcription of the singed locus of Drosophila melanogaster. Genetics 129:10731084.
150. Polard, P.,, and M. Chandler. 1995. Bacterial transposases and retroviral integrases. Mol. Microbiol. 15:1323.
151. Powers, T.,, and P. Walter. 1995. Reciprocal stimulation of GTP hydrolysis by two directly interacting GTPases. Science 269:14221424.
152. Preston, C. R.,, and W. R. Engels. 1996. P-element-induced male recombination and gene conversion in Drosophila. Genetics 144:16111622.
153. Preston, C. R.,, J. A. Sved,, and W. R. Engels. 1996. Flanking duplications and deletions associated with P-induced male recombination in Drosophila. Genetics 144:16231638.
154. Ramsden, D. A.,, D. C. van Gent,, and M. Gellert. 1997. Specificity in V(D)J recombination: new lessons from biochemistry and genetics. Curr. Opin. Immunol. 9:114120.
155. Rasmusson, K. E.,, J. D. Raymond,, and M. J. Simmons. 1993. Repression of hybrid dysgenesis in Drosophila melanogaster by individual naturally occurring P elements. Genetics 133: 605622.
156. Rasmusson, K. E.,, M. J. Simmons,, J. D. Raymond,, and C. F. McLarnon. 1990. Quantitative effects of P elements on hybrid dysgenesis in Drosophila melanogaster. Genetics 124: 647662.
157. Reed, R. 1996. Initial splice-site recognition and pairing during pre-mRNA splicing. Curr. Opin. Genet. Dev. 6:215220.
158. Rice, P.,, R. Craigie,, and D. R. Davies. 1996. Retroviral integrases and their cousins. Curr. Opin. Struct. Biol. 6:7683.
159. Rio, D. C. 1990. Molecular mechanisms regulating Drosophila P element transposition. Annu. Rev. Genet. 24: 543578.
160. Rio, D. C. 1991. Regulation of Drosophila P element transposition. Trends Genet. 7:282287.
161. Rio, D. C.,, F. A. Laski,, and G. M. Rubin. 1986. Identification and immunochemical analysis of biologically active Drosophila P element transposase. Cell 44:2132.
162. Rio, D. C.,, and G. M. Rubin. 1988. Identification and purification of a Drosophila protein that binds to the terminal 31- base-pair inverted repeats of the P transposable element. Proc. Natl. Acad. Sci. USA 85:89298933.
163. Robertson, H. M.,, and W. R. Engels. 1989. Modified P elements that mimic the P cytotype in Drosophila melanogaster. Genetics 123:815824.
164. Robertson, H. M.,, C. R. Preston,, R. W. Phillis,, D. M. Johnson- Schlitz,, W. K. Benz,, and W. R. Engels. 1988. A stable genomic source of P element transposase in Drosophila melanogaster. Genetics 118:461470.
165. Roche, S. E.,, and D. C. Rio. 1998. Trans-silencing by P elements inserted in subtelomeric heterochromatin involves the Drosophila Polycomb group gene, Enhancer of zeste. Genetics 149:18391855.
166. Roche, S. E.,, M. Schiff,, and D. C. Rio. 1995. P-element repressor autoregulation involves germ-line transcriptional repression and reduction of third intron splicing. Genes Dev. 9:12781288.
167. Roiha, H.,, G. M. Rubin,, and K. O’Hare. 1988. P element insertions and rearrangements at the singed locus of Drosophila melanogaster. Genetics 119:7583.
168. Rong, Y. S.,, and K. G. Golic. 2000. Gene targeting by homologous recombination in drosophila. Science 228:20132018.
169. Ronsseray, S.,, M. Lehmann,, and D. Anxolabehere. 1991. The maternally inherited regulation of P elements in Drosophila melanogaster can be elicited by two P copies at cytological site 1A on the X chromosome. Genetics 129:501512.
170. Ronsseray, S.,, M. Lehmann,, D. Nouaud,, and D. Anxolabehere. 1996. The regulatory properties of autonomous subtelomeric P elements are sensitive to a suppressor of variegation in Drosophila melanogaster. Genetics 143:16631674. (Erratum, 144:1329.)
171. Ronsseray, S.,, B. Lemaitre,, and D. Coen. 1993. Maternal inheritance of P cytotype in Drosophila melanogaster: a ‘‘pre- P cytotype’’ is strictly extra-chromosomally transmitted. Mol. Gen. Genet. 241:115123.
172. Ronsseray, S.,, L. Marin,, M. Lehmann,, and D. Anxolabehere. 1998. Repression of hybrid dysgenesis in Drosophila melanogaster by combinations of telomeric P-element reporters and naturally occurring P elements. Genetics 149:18571866.
173. Rørth, P. 1996. A modular misexpression screen in Drosophila detecting tissue-specific phenotypes. Proc. Natl. Acad. Sci. USA 93:1241812422.
174. Rørth, P. 1998. Gal4 in the Drosophila female germline. Mech. Dev. 78:113118.
175. Rørth, P.,, K. Szabo,, A. Bailey,, T. Laverty,, J. Rehm,, G. M. Rubin,, K. Weigmann,, M. Milan,, V. Benes,, and W. Ansorge. 1998. Systematic gain-of-function genetics in Drosophila. Development 125:10491057.
176. Roth, D. B.,, and M. Gellert. 2000. New guardians of the genome. Nature 404:823825.
177. Rubin, G. M.,, L. Hong,, P. Brokstein,, M. Evans-Holm,, E. Frise,, M. Stapleton,, and D. A. Harvey. 2000. A Drosophila complementary DNA resource. Science 287:22222224.
178. Rubin, G. M.,, M. G. Kidwell,, and P. M. Bingham. 1982. The molecular basis of P-M hybrid dysgenesis: the nature of induced mutations. Cell 29:987994.
179. Rubin, G. M.,, and E. B. Lewis. 2000.Abrief history of Drosophila’s contributions to genome research. Science 287: 22162218.
180. Rubin, G. M.,, and A. C. Spradling. 1982. Genetic transformation of Drosophila with transposable element vectors. Science 218:348353.
181. Rubin, G. M.,, M. D. Yandell,, J. R. Wortman,, G. L. Gabor Miklos,, C. R. Nelson,, I. K. Hariharan,, M. E. Fortini,, P. W. Li,, R. Apweiler,, W. Fleischmann,, J. M. Cherry,, S. Henikoff,, M. P. Skupski,, S. Misra,, M. Ashburner,, E. Birney,, M. S. Boguski,, T. Brody,, P. Brokstein,, S. E. Celniker,, S. A. Chervitz,, D. Coates,, A. Cravchik,, A. Gabrielian,, R. F. Galle,, W. M. Gelbart,, R. A. George,, L. S. Goldstein,, F. Gong,, P. Guan,, N. L. Harris,, B. A. Hay,, R. A. Hoskins,, J. Li,, Z. Li,, R. O. Hynes,, S. J. Jones,, P. M. Kuehl,, B. Lemaitre,, J. T. Littleton,, D. K. Morrison,, C. Mungall,, P. H. O’Farrell,, O. K. Pickeral,, C. Shue,, L. B. Vosshall,, J. Zhang,, Q. Zhao,, X. H. Zheng,, F. Zhong,, W. Zhong,, R. Gibbs,, J. C. Venter,, M. D. Adams,, and S. Lewis. 2000. Comparative genomics of the eukaryotes. Science 287:22042215.
182. Savilahti, H.,, P. A. Rice,, and K. Mizuuchi. 1995. The phage Mu transpososome core: DNA requirements for assembly and function. EMBO J. 14:48934903.
183. Schmid, S. L.,, M. A. McNiven,, and P. De Camilli. 1998. Dynamin and its partners: a progress report. Curr. Opin. Cell. Biol. 10:504512.
184. Scottoline, B. P.,, S. Chow,, V. Ellison,, and P. O. Brown. 1997. Disruption of the terminal base pairs of retroviral DNA during integration. Genes Dev. 11:371382.
185. Sentry, J. W.,, and K. Kaiser. 1994. Application of inverse PCR to site-selected mutagenesis of Drosophila. Nucleic Acids Res. 22:34293430.
186. Sepp, K. J.,, and V. J. Auld. 1999. Conversion of lacZ enhancer trap lines to GAL4 lines using targeted transposition in Drosophila melanogaster. Genetics 151:10931101.
187. Sharp, P. A. 1999. RNAi and double-strand RNA. Genes Dev. 13:139141.
188. Siebel, C. W.,, A. Admon,, and D. C. Rio. 1995. Soma-specific expression and cloning of PSI, a negative regulator of P element pre-mRNA splicing. Genes Dev. 9:269283.
189. Siebel, C. W.,, L. D. Fresco,, and D. C. Rio. 1992. The mechanism of somatic inhibition of Drosophila P element premRNA splicing: multiprotein complexes at an exon pseudo- 5′ splice site control U1 snRNP binding. 6:13861401.
190. Siebel, C. W.,, R. Kanaar,, and D. C. Rio. 1994. Regulation of tissue-specific P-element pre-mRNA splicing requires the RNA-binding protein PSI. Genes Dev. 8:17131725.
191. Siebel, C. W.,, and D. C. Rio. 1990. Regulated splicing of the Drosophila P transposable element third intron in vitro: somatic repression. Science 248:12001208.
192. Simmons, M. J.,, and L. M. Bucholz. 1985. Transposase titration in Drosophila melanogaster: a model of cytotype in the P-M system of hybrid dysgenesis. Proc. Natl. Acad. Sci. USA 82:81198123.
193. Simmons, M. J.,, J. D. Raymond,, C. D. Grimes,, C. Belinco,, B. C. Haake,, M. Jordan,, C. Lund,, T. A. Ojala,, and D. Papermaster. 1996. Repression of hybrid dysgenesis in Drosophila melanogaster by heat- shock-inducible sense and antisense P-element constructs. Genetics 144:15291544.
194. Simmons, M. J.,, J. D. Raymond,, K. E. Rasmusson,, L. M. Miller,, C. F. McLarnon,, and J. R. Zunt. 1990. Repression of P element-mediated hybrid dysgenesis in Drosophila melanogaster. Genetics 124:663676.
195. Smith, C. W.,, and J. Valcarcel. 2000. Alternative pre-mRNA splicing: the logic of combinational control. Trends Biochem. Sci. 25:381388.
196. Smith, G. C.,, R. B. Cary,, N. D. Lakin,, B. C. Hann,, S. H. Teo,, D. J. Chen,, and S. P. Jackson. 1999. Purification and DNA binding properties of the ataxia-telangiectasia gene product ATM. Proc. Natl. Acad. Sci. USA 96:1113411139.
197. Smith, G. C.,, and S. P. Jackson. 1999. The DNA-dependent protein kinase. Genes Dev. 13:916934.
198. Spradling, A. C. 1986. P element-mediated transformation, p. 175197. In Drosophila: A Practical Approach. RL Press, Oxford, England.
199. Spradling, A. C.,, and G. M. Rubin. 1982. Transposition of cloned P elements into Drosophila germ line chromosomes. Science 218:341347.
200. Spradling, A. C.,, D. Stern,, E. J. Rhem,, T. Laverty,, N. Mozden,, S. Misra,, and G. M. Rubin. 1999. The Berkeley Drosophila Genome Project gene disruption project: Single P-element insertions mutating 25% of vital Drosophila genes. Genetics 153:135177.
201. Spradling, A. C.,, D. M. Stern,, I. Kiss,, J. Roote,, T. Laverty,, and G. M. Rubin. 1995. Gene disruptions using P transposable elements: an integral component of the Drosophila genome project. Proc. Natl. Acad. Sci. USA 92:1082410830.
202. Staveley, B. E.,, T. R. Heslip,, R. B. Hodgetts,, and J. B. Bell. 1995. Protected P-element termini suggest a role for invertedrepeat- binding protein in transposase-induced gap repair in Drosophila melanogaster. Genetics 139:13211329.
203. Stellwagen, A. E.,, and N. L. Craig. 1997. Gain-of-function mutations in TnsC, an ATP-dependent transposition protein that activates the bacterial transposon Tn7. Genetics 145: 573585.
204. Stellwagen, A. E.,, and N. L. Craig. 1998. Mobile DNA elements: controlling transposition with ATP-dependent molecular switches. Trends Biochem. Sci. 23:486490.
205. Struhl, G.,, and K. Basler. 1993. Organizing activity of wingless protein in Drosophila. Cell 72:527540.
206. Sved, J. A.,, L. M. Blackman,, A. S. Gilchrist,, and W. R. Engels. 1991. High levels of recombination induced by homologous P elements in Drosophila melanogaster. Mol. Gen. Genet. 225:443447.
207. Sved, J. A.,, L. M. Blackman,, Y. Svoboda,, and R. Colless. 1995. Male recombination with single and homologous P elements in Drosophila melanogaster. Mol. Gen. Genet. 246: 381386.
208. Sved, J. A.,, W. B. Eggleston,, and W. R. Engels. 1990. Germline and somatic recombination induced by in vitro modified P elements in Drosophila melanogaster. Genetics 124: 331337.
209. Svoboda, Y. H.,, M. K. Robson,, and J. A. Sved. 1995. Pelement- induced male recombination can be produced in Drosophila melanogaster by combining end-deficient elements in trans. Genetics 139:16011610.
210. Tanaka, M. M.,, X. M. Liang,, Y. H. Gray,, and J. A. Sved. 1997. The accumulation of P-element-induced recombinants in the germline of male Drosophila melanogaster. Genetics 147:17691782.
211. Theodosiou, N. A.,, and T. Xu. 1998. Use of FLP/FRT system to study Drosophila development. Methods 14:355365.
212. Thompson-Stewart, D.,, G. H. Karpen,, and A. C. Spradling. 1994. A transposable element can drive the concerted evolution of tandemly repetitious DNA. Proc. Natl. Acad. Sci. USA 91:90429046.
213. Tower, J.,, G. H. Karpen,, N. Craig,, and A. C. Spradling. 1993. Preferential transposition of Drosophila P elements to nearby chromosomal sites. Genetics 133:347359.
214. Tseng, J. C.,, S. Zollman,, A. C. Chain,, and F. A. Laski. 1991. Splicing of the Drosophila P element ORF2-ORF3 intron is inhibited in a human cell extract. Mech. Dev. 35:6572.
215. Vos, J. C.,, I. De Baere,, and R. H. Plasterk. 1996. Transposase is the only nematode protein required for in vitro transposition of Tc1. Genes Dev. 10:755761.
216. Wang, J. C. 1996. DNA topoisomerases. Annu. Rev. Biochem. 65:635692.
217. Weaver, D. T. 1995. What to do at an end: DNA doublestrand- break repair. Trends Genet. 11:388392.
218. Wilson, C.,, R. K. Pearson,, H. J. Bellen,, C. J. O’Kane,, U. Grossniklaus,, and W. J. Gehring. 1989. P-element-mediated enhancer detection: an efficient method for isolating and characterizing developmentally regulated genes in Drosophila. Genes Dev. 3:13011313.
219. Wittinghofer, A.,, and E. F. Pai. 1991. The structure of Ras protein: a model for a universal molecular switch. Trends Biochem. Sci. 16:382387.
220. Xu, T.,, and S. D. Harrison. 1994. Mosaic analysis using FLP recombinase. Methods Cell Biol. 44:655681.
221. Xu, T.,, and G. M. Rubin. 1993. Analysis of genetic mosaics in developing and adult Drosophila tissues. Development 117: 12231237.
222. Yamauchi, M.,, and T. A. Baker. 1998. An ATP-ADP switch in MuB controls progression of the Mu transposition pathway. EMBO J. 17:55095518.
223. Zhang, P.,, and A. C. Spradling. 1993. Efficient and dispersed local P element transposition from Drosophila females. Genetics 133:361373.

Tables

Generic image for table
Table 1

Genetic assays for P cytotype repression

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21
Generic image for table
Table 2

Sequences of 5′ splice sites

Citation: Rio D. 2002. P Transposable Elements in Drosophila melanogaster, p 484-518. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch21

This is a required field
Please enter a valid email address
Please check the format of the address you have entered.
Approval was a Success
Invalid data
An Error Occurred
Approval was partially successful, following selected items could not be processed due to error