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There are many helicases resulting from the great variety of processes in which strand separation must be catalyzed. Helicases are often used to separate strands of a DNA double helix or a self-annealed RNA molecule using the energy from ATP hydrolysis, a process characterized by the breaking of hydrogen bonds between annealed nucleotide bases.

They also function to remove nucleic acid-associated proteins and catalyze homologous DNA recombination.

Helicases adopt different structures and oligomerization states. Whereas DnaB -like helicases unwind DNA as ring-shaped hexamers , other enzymes have been shown to be active as monomers or dimers.

Studies have shown that helicases may act passively, waiting for uncatalyzed unwinding to take place and then translocating between displaced strands, [4] or can play an active role in catalyzing strand separation using the energy generated in ATP hydrolysis.

Factors that contribute to the height of the activation barrier include: The size of the activation barrier to overcome by the helicase contributes to its classification as an active or passive helicase.

These two categories of helicases may also be modelled as mechanisms. In such models the passive helicases are conceptualized as Brownian ratchets, driven by thermal fluctuations and subsequent anisotropic gradients across the DNA lattice.

The active helicases, in contrast, are conceptualized as stepping motors - also known as powerstroke motors - utilizing either a conformational "inch worm" or a hand-over-hand "walking" mechanism to progress.

DNA helicases were discovered in E. The common function of helicases accounts for the fact that they display a certain degree of amino acid sequence homology ; they all possess sequence motifs located in the interior of their primary structure , involved in ATP binding, ATP hydrolysis and translocation along the nucleic acid substrate.

The variable portion of the amino acid sequence is related to the specific features of each helicase. The presence of these helicase motifs allows putative helicase activity to be attributed to a given protein, but does not necessarily confirm it as an active helicase.

Conserved motifs do, however, support an evolutionary homology among enzymes. Based on these helicase motifs, a number of helicase superfamilies have been distinguished.

Helicases are classified in 6 groups superfamilies based on their shared sequence motifs. They are also classified by translocation polarity.

Various types of mutations found in ATRX have been found to be associated with ATR-X, including most commonly single-base missense mutations, as well as nonsense, frameshift, and deletion mutations.

XPD helicase mutations leading to trichothiodystrophy are found throughout the protein in various locations involved in protein-protein interactions.

It has been suggested that XPD helicase mutations leading to Cockayne syndrome could be the result of mutations within XPD, causing rigidity of the protein and subsequent inability to switch from repair functions to transcription functions due to a "locking" in repair mode.

Bloom syndrome is characterized by a predisposition to cancer with early onset, with a mean age-of-onset of 24 years. Werner syndrome is a disorder of premature aging, with symptoms including early onset of atherosclerosis and osteoporosis and other age related diseases, a high occurrence of sarcoma, and death often occurring from myocardial infarction or cancer in the 4th to 6th decade of life.

Rothmund-Thomson syndrome, also known as poikiloderma congenitale , is characterized by premature aging, skin and skeletal abnormalities, rash, poikiloderma , juvenile cataracts, and a predisposition to cancers such as osteosarcomas.

During meiosis DNA double-strand breaks and other DNA damages in a chromatid are repaired by homologous recombination using either the sister chromatid or a homologous non-sister chromatid as template.

It was suggested that COs are restricted because of the long term costs of CO recombination, that is, the breaking up of favorable genetic combinations of alleles built up by past natural selection.

They also play an important role in sensing viral RNAs. On the other hand, ring-forming RNA helicases have been found in bacteria and viruses.

Although these proteins carry the typical helicase motifs, hydrolize ATP in a nucleic acid-dependent manner, and are built around a helicase core, in general, no unwinding activity is observed.

RNA helicases that do exhibit unwinding activity have been characterized by at least two different mechanisms: Canonical duplex unwinding is the stepwise directional separation of a duplex strand, as described above, for DNA unwinding.

However, local strand separation occurs by a process wherein the helicase enzyme is loaded at any place along the duplex.

This is usually aided by a single-strand region of the RNA, and the loading of the enzyme is accompanied with ATP binding.

An RNA helicase database [62] is currently available online that contains a comprehensive list of RNA helicases with information such as sequence, structure, and biochemical and cellular functions.

Various methods are used to measure helicase activity in vitro. These methods range from assays that are qualitative assays that usually entail results that do not involve values or measurements to quantitative assays with numerical results that can be utilized in statistical and numerical analysis.

In , the first direct biochemical assay was developed for measuring helicase activity. Other methods were later developed that incorporated some, if not all of the following: Commercially available diagnostic kits are also available.

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