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The Tn21 subgroup of bacterial transposable elements.

The Tn3 family of transposable elements is probably the most successful group of mobile DNA elements in bacteria: there are many different but related members and they are widely distributed in gram-negative and gram-positive bacteria. The Tn21 subgroup of the Tn3 family contains closely related elements that provide most of the currently known variation in Tn3-like elements in gram-negative bacteria and that are largely responsible for the problem of multiple resistance to antibiotics in these organisms. This paper reviews the structure, the mechanism of transposition, the mode of acquisition of accessory genes, and the evolution of these elements.

Bacteria↗

Survey of transposable elements from rice genomic sequences.

Oryza sativa L. (domesticated rice) is a monocotyledonous plant, and its 430 Mb genome has been targeted for complete sequencing. We performed a high-resolution computer-based survey for transposable elements on 910 Kb of rice genomic DNA sequences. Both class I and II transposable elements were present, contributing 19.9% of the sequences surveyed. Class II elements greatly outnumbered class I elements (166 versus 22), although class I elements made up a greater percentage (12.2% versus 6.6%) of nucleotides surveyed. Several Mutator-like elements (MULEs) were identified, including rice elements that harbor truncated host cellular genes. MITEs (miniature inverted-repeat transposable elements) account for 71.6% of the mined transposable elements and are clearly the predominant type of transposable element in the sequences examined. Moreover, a putative Stowaway transposase has been identified based on shared sequence similarity with the mined MITEs and previously identified plant mariner-like elements (MLEs). Members of a group of novel rice elements resembling the structurally unusual members of the Basho family in Arabidopsis suggest a wide distribution of these transposons among plants. Our survey provides a preview of transposable element diversity and abundance in rice, and allows for comparison with genomes of other plant species.

Base Sequence↗

Constitutive heterochromatin and transposable elements in Drosophila melanogaster.

Several families of transposable elements (TEs), most of them belonging to the retrotransposon category, are particularly enriched in Drosophila melanogaster constitutive heterochromatin. The enrichment of TE-homologous sequences into heterochromatin is not a peculiar feature of the Drosophila genome, but appears to be widespread among higher eukaryotes. The constitutive heterochromatin of D. melanogaster contains several genetically active domains; this raises the possibility that TE-homologous sequences inserted into functional heterochromatin compartments may be expressed. In this review, I present available data on the genetic and molecular organization of D. melanogaster constitutive heterochromatin and its relationship with transposable elements. The implications of these findings on the possible impact of heterochromatic TEs on the function and evolution of the host genome are also discussed.

Animals↗

Tourist C transposable elements are closely associated with genes expressed in flowers of rice (Oryza sativa).

Tourist elements comprise a group of transposable elements in plants. One of these elements, Tourist-OsaCatA(a Tourist C element), has been found in the 5; flanking region of a catalase gene, CatA, in rice (Oryza sativa). Using reverse transcriptase-PCR (RT-PCR) analyses of leaves, roots, flowers and developing seeds of rice, we assessed the transcription levels of ten known genes containing Tourist C elements, and of three additional putative genes for which expressed sequence tags (ESTs) including Tourist C elements have been isolated. We found that nine of the ten known genes and two of the three represented by ESTs were expressed in at least one of the organs we analyzed, and all of the genes detected were expressed in flowers, usually in stamens or pistils. We also assessed the expression of the 29 Tourist C-containing hypothetical coding sequences (CDSs) obtained so far by high-throughput genomic sequencing. We found that CDSs of all 11 genes whose transcripts were detectable by RT-PCR were expressed in flowers, especially in stamens or pistils. In contrast, RT-PCR analyses of genes or CDSs associated with other miniature inverted-repeat transposable elements (MITEs), such as Tourist D, Gaijin, Explorer, and Castaway, showed that some of them were expressed only minimally or not at all in flowers. Therefore, compared with other MITEs, Tourist C elements seem to show a strong association with genes that are expressed in the flowers of rice.

DNA Transposable Elements↗

Molecular and functional characterization of Slide, an Ac-like autonomous transposable element from tobacco.

A new transposable element of tobacco, Slide, was isolated from the tl mutant line, which shows somatic instability, after its transposition into a locus encoding nitrate reductase (NR). The Slide-124 element is 3733 bp long and its coding sequences show similarities with conserved domains of the transposases of Ac, Tam3 and hobo. Excision from the NR locus is detectable in somatic leaf tissues and Slide mobility is triggered by in vitro tissue culture. Slide excision events create footprints similar to those left by Ac and Tam3. Tobacco lines derived from the tl mutant line seem characterized by unmethylated copies of a few members of the highly repetitive Slide family. Slide mobility was monitored in transient expression assays. In wild-type tobacco protoplasts, the complete Slide element, as well as a defective copy, is able to excise. The complete Slide element, but not the defective version, is able to excise in protoplasts of the heterologous species lettuce (Lactuca sativa). These results show that Slide carries the functions required for its own mobility, and represents the first autonomous Ac-like element characterized in Solanaceae species.

Amino Acid Sequence↗

Flipper, a mobile Fot1-like transposable element in Botrytis cinerea.

A transposable element, Flipper, was isolated from the phytopathogenic fungus Botrytis cinerea. The element was identified as an insertion sequence within the coding region of the nitrate reductase gene. The Flipper sequence is 1842 bp long with perfect inverted terminal repeats (ITRs) of 48 bp and an open reading frame (ORF) of 533 amino acids, potentially encoding for a transposase; the element is flanked by the dinucleotide TA. The encoded protein is very similar to the putative transposases of three elements from other phytopathogenic fungi, Fot1 from Fusarium oxysporum, and Pot2 and MGR586 from Magnaporthe grisea. The number of Flipper elements in strains of B. cinerea varied from 0 to 20 copies per genome. Analysis of the descendants of one cross showed that the segregation ratio of Flipper elements was 2:2 and that the copies were not linked.

Amino Acid Sequence↗

The Queensland fruit fly, Bactrocera tryoni, contains multiple members of the hAT family of transposable elements.

Members of the hAT transposable element family are mobile in non-host insect species and have been used as transformation vectors in some of these species. We report that the Queensland fruit fly, Bactrocera tryoni, contains at least two types of insect hAT elements called Homer and a Homer-like element (HLE). The Homer element is 3789 bp in size and contains 12-bp imperfect inverted terminal repeats. The Homer element contains a long open reading frame (ORF) that encodes a putative transposase. Three different copies of this long ORF were recovered from the B. tryoni genome and, upon transcription and translation in an in vitro system, all produced transposase. The HLE is an incomplete element since no 3' inverted terminal repeat (ITR) was found. Homer and the HLE are as related to one another as either is to the other insect hAT elements such as Hermes, hobo, hermit and hopper. The structure and distribution of these two Homer elements is described.

Amino Acid Sequence↗

Analysis of a transposable element in Caenorhabditis elegans.

A transposable element, designated Tc1, has been characterized in Caenorhabditis elegans. Tc1 is 1.7 kilobases long, has an inverted terminal repeat of less than 100 base pairs, and is repeated as a highly conserved element. The copy number and genomic positions of Tc1 are extremely variable among strains, implying that Tc1 is mobile. However, progeny of interstrain crosses did not show hybrid dysgenic traits that might be due to Tc1 transposition.

Animals↗

The isolation of Ant1, a transposable element from Aspergillus niger.

A transposable element has been isolated from the industrially important fungus Aspergillus niger (strain N402). The element was identified as an insertion sequence within the coding region of the nitrate reductase gene. It had inserted at a TA site and appeared to have duplicated the target site upon insertion. The isolated element was found to be 4798 bp in length and contained 37-bp inverted, imperfect, terminal repeats (ITRs). The sequence of the central region of the element revealed an open reading frame (designated ORF1) which showed similarity, at the amino acid level, to the transposase of the Tc1/mariner class of DNA transposons. Another sequence within the central region of the element showed similarity to the 3' coding and downstream untranslated region of the amyA gene of A. niger. Sequence homology and structural features indicate that this element, which has been named Ant1 (A. niger transposon 1), is related to the Tc1/mariner group of DNA transposons. Ant1 is apparently present as a single copy in strain N402 of A. niger.

Amino Acid Sequence↗

A new family of the poly-deoxyadenylated class of Drosophila transposable elements identified by a representative member at the dunce locus.

Two transposable elements have been identified at the dunce locus in chromosomes recovered after a premeiotic and interchromosomal conversion event occurred at this gene. One is approximately 8.2 kb and is inserted near the 5' end of the gene. This element was identified by sequence analysis as a member of the B104 (roo) family of copia-like transposable elements. The second resides near the 3' end of the gene and represents a new family of the class of poly-deoxyadenylated [poly(dA)] transposable elements. It is 0.38 kb in length and has one terminus consisting of a stretch of 29 deoxyadenosine residues with a polyadenylation site like those found in mRNA molecules, located about 20 bp away from the poly(dA) stretch. Fourteen base pairs of genome DNA is duplicated at the target site of this element.

3',5'-Cyclic-AMP Phosphodiesterases↗

[Transposable elements in the Animal Kingdom].

Transposable elements (TEs) are commonly thought to be of universal occurrence in eukaryotes. Analysis of complete higher eukaryotic genomes confirms TE status as substantial genome components and provides insights into their role in shaping the genome structure of extant eukaryotes. This review addresses several recently investigated problems in transposon biology, including potential roles of promoter organization in transposon function and evolution, the ubiquity of TEs in numerous phyla of the animal kingdom, and possible connections between transposon content and the mode of reproduction.

Animals↗

The distribution of transposable elements on X chromosomes from a natural population of Drosophila simulans.

The distribution of 13 transposable element families along 15 X chromosomes from an African natural population of Drosophila simulans was determined by in situ hybridization to polytene chromosomes. The transposable elements cloned from Drosophila melanogaster all hybridized with Drosophila simulans chromosomes. The number of copies per family was 3.5 times lower in the latter species and correlated with the copy number per family in Drosophila melanogaster. With the exception of 297, the copy number per chromosome followed a Poisson distribution. Element frequencies per chromosome band were generally low. However, several sites of the distal region and the base of the X chromosome had high frequencies of occupation. Elements had higher abundance at the base of the chromosome compared to distal regions. Overall, the distribution of transposable elements in Drosophila simulans is similar to that found in Drosophila melanogaster. These data provide evidence for the operation of a force (or forces) opposing transpositional increase in copy number, and that this force is weaker at the bases of chromosomes, consistent with the idea that recombination between elements at non-homologous sites contains TE copy number. The reduction in copy number of all TE families in Drosophila simulans compared to Drosophila melanogaster can be explained by stronger selection against transposable element multiplication and/or lower rates of transposition in Drosophila simulans.

Africa↗

Insertion and excision of the transposable element mariner in Drosophila.

The transposable element mariner is active in both germline and somatic cells of Drosophila mauritiana. Activity of the element is greatly enhanced in the presence of Mos1, a genetic factor identified as an autonomous copy of mariner. A strain of D. mauritiana containing Mos1 and other copies of mariner was used to initiate a screen for visible mutations. More than 20 mutations were obtained, including alleles of white, yellow and vermilion. Six alleles were characterized at the molecular level, and all were found to contain a mariner element inserted into the affected gene. Four insertions into the white locus were sequenced to determine the exact site of insertion of mariner. There appears to be little sequence specificity requirement for mariner insertion, other than an absolute requirement for the dinucleotide TA, which is duplicated upon insertion. Sequences of phenotypically wild-type germline and somatic revertants obtained from various white alleles, including the previously isolated wpch allele, were obtained using the polymerase chain reaction. Mariner excision is imprecise in both germline and soma, and the most frequent excision events are the same in the two tissues. Mutant derivatives of wpch were also studied, and were found to exhibit a wide range of molecular structures and phenotypes.

Alleles↗

Invasion of Drosophila virilis by the Penelope transposable element.

The Penelope family of transposable elements (TEs) is broadly distributed in most species of the virilis species group of Drosophila. This element plays a pivotal role in hybrid dysgenesis in Drosophila virilis, in which at least four additional TE families are also activated. Here we present evidence that the Penelope family of elements has recently invaded D. virilis. This evidence includes: (1) a patchy geographical distribution, (2) genomic locations mainly restricted to euchromatic chromosome arms in various geographical strains, and (3) a high level of nucleotide similarity among members of the family. Two samples from a Tashkent (Middle Asia) population of D. virilis provide further support for the invasion hypothesis. The 1968 Tashkent strain is free of Penelope sequences, but all individuals collected from a 1997 population carry at least five Penelope copies. Furthermore, a second TE, Ulysses, has amplified and spread in this population. These results provide evidence for the Penelope invasion of a D. virilis natural population and the mobilization of unrelated resident transposons following the invasion.

Animals↗

Effects of recombination rate and gene density on transposable element distributions in Arabidopsis thaliana.

Transposable elements (TEs) comprise a major component of eukaryotic genomes, and exhibit striking deviations from random distribution across the genomes studied, including humans, flies, nematodes, and plants. Although considerable progress has been made in documenting these patterns, the causes are subject to debate. Here, we use the genome sequence of Arabidopsis thaliana to test for the importance of competing models of natural selection against TE insertions. We show that, despite TE accumulation near the centromeres, recombination does not generally correlate with TE abundance, suggesting that selection against ectopic recombination does not influence TE distribution in A. thaliana. In contrast, a consistent negative correlation between gene density and TE abundance, and a strong under-representation of TE insertions in introns suggest that selection against TE disruption of gene expression is playing a more important role in A. thaliana. High rates of self-fertilization may reduce the importance of recombination rate in genome structuring in inbreeding organisms such as A. thaliana and Caenorhabditis elegans.

Arabidopsis↗

On two transposable elements from Bacillus stearothermophilus.

Two transposable elements, IS5376 and IS5377, were identified in the thermophile Bacillus stearothermophilus CU21 based upon the following criteria: (1) both were found to appear on different plasmids introduced into the same host CU21; (2) signals of homology were found between the genomic DNA of CU21 and each of them; (3) different numbers of Southern hybridization bands were found for the genomic DNA of different strains of B. stearothermophilus; and (4) characteristic inverted repeats at both ends and direct repeats of the target DNA adjacent to them were found for both IS5376 and IS5377. Two open reading frames (ORFs) were detected for IS5376 and one for IS5377. The putative coding products of the ORFs are homologous to those of known ISs from mesophiles and are considered to be transposases. The results of analyses of nucleotide sequence and the deduced amino acid sequence suggest that IS5376 is a member of the IS21 family and that IS5377 is a member of the IS4 family.

Amino Acid Sequence↗

Characterization and zeste reaction of spontaneously reduced transposable elements in Drosophila melanogaster.

Seven transposable elements (TE), selected from parental stock w 239 Cy, have been characterized. They all carry w+ but have lost both rst+ and the two polytene chromosome bands of a complete TE and are thus cytologically invisible. In situ hybridization with probes from the white-roughest interval shows that the reduced TE's are truncated in the same region, 40-90 kb upstream of the white locus, well proximal to the zeste interaction site. They express the white gene in wild type fashion in all tissues studied (eyes, testis sheaths, and Malpighian tubules) but give zeste reactions varying from yellow to bright red. Thus, by the criterion of the zeste reaction, 40 kb of TE sequences 5' to the transcription start site cannot insulate the white gene from position effects. One single spontaneously reduced TE (TE203) originated from parental stock w TE100 ct. This TE is truncated closer to the white locus (+ 18- +26 kb), but still well separated from the zeste interaction site. In a z1w11E4 background, homozygous TE203 flies have an uneven eye pigmentation, while heterozygotes have eyes that are yellowish red. The Malpighian tubules and testes show wild type pigmentation in both cases, like normal TE's.

Animals↗

How valuable are model organisms for transposable element studies?

Model organisms have proved to be highly informative for many types of genetic studies involving 'conventional' genes. The results have often been successfully generalized to other closely related organisms and also, perhaps surprisingly frequently, to more distantly related organisms. Because of the wealth of previous knowledge and their availability and convenience, model organisms were often the species of choice for many of the earlier studies of transposable elements. The question arises whether the results of genetic studies of transposable elements in model organisms can be extrapolated in the same ways as those of conventional genes? A number of observations suggest that special care needs to be taken in generalizing the results from model organisms to other species. A hallmark of many transposable elements is their ability to amplify rapidly in species genomes. Rapid spread of a newly invaded element throughout a species range has also been demonstrated. The types and genomic copy numbers of transposable elements have been shown to differ greatly between some closely related species. Horizontal transfer of transposable elements appears to be more frequent than for nonmobile genes. Furthermore, the population structure of some model organisms has been subject to drastic recent changes that may have some bearing on their transposable element genomic complements. In order to initiate discussion of this question, several case studies of transposable elements in well-studied Drosophila species are presented.

Biological Evolution↗