INTRODUCTION:

Transposons are sequences of DNA that can move around to different positions within the genome of a single cell, a process called transposition. In the process, they can cause mutations and change the amount of DNA in the genome. Transposons were also once called jumping genes, and are examples of mobile genetic elements. They were discovered by Barbara McClintock. Transposons make up a large fraction of genome sizes which is evident through the Cvalues of eukaryotic species. The sheer volume of seemingly useless material initially puzzled researchers, so that it was termed "Junk DNA" until further research revealed the critical role that it played in the development of an organism. They are very useful to researchers as a means to alter DNA inside a living organism.

TYPES OF TRANSPOSONS:
There are many different types of Transposon Elements(TEs), as well as a number of ways to categorize them. One of the more common divisions is between those TEs that require reverse transcription (i.e., the transcription of RNA intoDNA) in order to transpose and those that do not. The former elements are known as retrotransposons or class 1 TEs, whereas the latter are known as DNA transposons or class 2 TEs.

DNA TRANSPOSONS:
All complete or "autonomous" class 2 TEs encode the protein transposase, which they require for insertion and excision. Some of these TEs also encode other proteins.DNA transposons never use RNA intermediariesthey always move on their own, inserting or excising themselves from the genome by means of a so-called "cut and paste" mechanism.

Class 2 TEs are characterized by the presence of terminal inverted repeats, about 9 to 40 base pairs long, on both of their ends. As the name suggests, terminal inverted repeats are inverted complements of each other; for instance, the complement of ACGCTA (the inverted repeat on the right side of the TE in the figure) is TGCGAT (which is the reverse order of the terminal inverted repeat on the left side of the TE in the figure). One of the roles of terminal inverted repeatsis to be recognized by transposase.

Less than 2% of the human genome is composed of class 2 TEs. This means that the majority of the substantial portion of the human genome that is mobile consists of the other major class of TEsthe retrotransposon

RETROTRANSPOSONS:
Unlike class 2 elements, class 1 elementsalso known as retrotransposonsmove through the action of RNAintermediaries. In other words, class 1 TEs do not encode transposase; rather, they produce RNA transcripts and then rely upon reverse transcriptase enzymes to reverse transcribe the RNA sequences back into DNA, which is then inserted into the target site. There are two major types of class 1 TEs: LTR retrotransposons, which are characterized by the presence of long terminal repeats (LTRs) on both ends; and non-LTR TEs, which lack the repeats. Both the LINE1, or L1, and Alu genes represent families of non-LTR TEs. In humans, these non-LTR TEs are the only active class of transposons; LTR retrotransposons and DNA transposons are only ancient genomic relics and are not capable of jumpin.

All TEs in both class 1 and class 2 contain flanking direct repeats. Flanking direct repeats are not actually part of the transposable element; rather, they play a role in insertion of the TE. Moreover, after a TE is excised, these repeats are left behind as "footprints." Sometimes, these footprints alter gene expression (i.e., expression of the gene in which they have been left behind) even after their related TE has moved to another location on the genome. Both class 1 and class 2 TEs can be either autonomous or nonautonomous. Autonomous TEs can move on their own, while nonautonomous elements require the presence of other TEs in order to move. This is because nonautonomous elements lack the gene for the transposase or reverse transcriptasethat is needed for their transposition, so they must "borrow" these proteins from another element in order to move.

RATE OF TRANSPOSITION, INDUCTION AND DEFENCE:

One study estimated the rate of transposition of a particular retrotransposon, the Ty1 element in Saccharomyces cerevisiae. Using several assumptions, the rate of successful transposition event per single Ty1 element came out to be about once every few months to once every few years. Most TEs appear to be silent in other words, these elements do not produce a phenotypic effect, nor do they actively move around the genome. At least that has been the general scientific consensus. Some silenced TEs are inactive because they have mutations that affect their ability to move from one chromosomal location to another; others are perfectly intact and capable of moving but are kept inactive by epigenetic defense mechanisms such as DNA methylation, chromatin remodeling, and miRNAs. In human cells, small interfering RNAs (siRNAs), also known as RNAi, can prevent transposition. RNAi is a naturally occurring mechanism that eukaryotes often use to regulate gene expression. Some transposable elements contain heat-shock like promoters and their rate of transposition increases if the cell is subjected to stress, thus increasing the mutation rate under these conditions, which might be beneficial to the cell. Transposons are found in all major branches of life. They may or may not have originated in the last universal common ancestor, or arisen independently multiple times, or perhaps arisen once and then spread to other kingdoms by horizontal gene transfer
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. While transposons may confer some benefits on their hosts

TRANSPOSONS CAUSING DISEASES:

Transposons are mutagens. They can damage the genome of their host cell in different ways: A transposon or a retroposon that inserts itself into a functional gene will most likely disable

that gene. After a transposon leaves a gene, the resulting gap will probably not be repaired correctly. Multiple copies of the same sequence, such as Alu sequences can hinder

precise chromosomal pairing during mitosis and meiosis, resulting in unequal crossovers, one of the main reasons for chromosome duplication. Diseases that are often caused by transposons include hemophilia A and B, severe combined immunodeficiency, porphyria, predisposition to cancer, and Duchenne muscular dystrophy. Additionally, many transposons contain promoters which drive transcription of their own transposase. These promoters can cause aberrant expression of linked genes, causing disease or mutant phenotypes.

ROLE OF TRANSPOSONS IN EVOLUTION:

Not all transposon jumping results in deleterious effects. In fact, transposons can drive the evolution of genomes by facilitating the translocation of genomic sequences, the shuffling of exons, and the repair of double-stranded breaks. Insertions and transposition can also alter gene regulatory regions and phenotypes. The fact that transposable elements do not always excise perfectly and can take genomic sequences along for the ride has also resulted in a phenomenon scientists call exon shuffling. Exon shuffling results in the juxtaposition of two previously unrelated exons, usually by transposition, thereby potentially creating novel gene products. The ability of transposons to increase genetic diversity, together with the ability of the genome to inhibit most TE activity, results in a balance that makes transposable elements an important part of evolution and gene regulation in all organisms that carry these sequences.

APPLICATIONS:
The first transposon was discovered in the plant maize (Zea mays, corn species), and is named dissociator (Ds). Likewise, the first transposon to be molecularly isolated was from a plant (Snapdragon). Appropriately, transposons have been an especially useful tool in plant molecular biology. Researchers use transposons as a means of mutagenesis. In this context, a transposon jumps into a gene and produces a mutation. The presence of the transposon provides a straightforward means of identifying the mutant allele, relative to chemical mutagenesis methods. Sometimes the insertion of a transposon into a gene can disrupt that gene's function in a reversible manner, in a process called insertional mutagenesis; transposase-mediated excision of the transposon restores gene function. This produces plants in which neighboring cells have different genotypes. This feature allows researchers to distinguish between genes that must be present inside of a cell in order to function (cell-autonomous) and genes that produce observable effects in cells other than those where the gene is expressed. Transposons are also a widely used tool for mutagenesis of most experimentally tractable organisms. The Tc1/mariner-class transposons piggyBac and Sleeping Beauty are active in mammalian cells and are being investigated for use in human gene therapy.
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REFERENCES:
Kazazian, H. H., & Moran, J. V. The impact of L1 retrotransposons on the human genome. Nature Genetics Feschotte, C., et al. Plant transposable elements: Where genetics meets genomics. Nature Reviews Genetics Lewin B (2000). Genes VII. Oxford University Press Kidwell, M.G. (2005). "Transposable elements". in ed. T.R. Gregory.