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This implies that operation 2 can be used to rearrange sequences in a DNA
molecule, thus accomplishing gene unscrambling.
The above operations are similar to the the “splicing operation” introduced
by Head [3] and circular splicing and mixed splicing [4, 14-16, 21]. It was
subsequently shown that some of these models have the computational power
of a universal Turing machine [1, 13, 22]. (See Head et al. [5] for a review.)
The process of gene unscrambling entails a series of successive or possibly
simultaneous intra- and intermolecular homologous recombinations. This is
followed by excision of all sequences
τ s y τ e , where the sequence y is marked
by the presence of telomere addition sequences
τ s for telomere “start” (at its
5' end), and
τ e for telomere “end” (at its 3' end). Thus, from a long sequence
u τ s y τ e v , this step retains only
τ s y τ e in the macronucleus. Last, the enzyme
telomerase extends the length of the telomeric sequences (usually double-
stranded TTTTGGGG repeats in these organisms) from
τ s and
τ e to protect
the ends of the DNA molecule.
We now make the assumption that, either by a structural alignment of the
DNA or by other biochemical factors, the cell decides which sequences are
non-protein-coding (IESs) and which are ultimately protein coding (MDSs),
as well as which are the pointers x . Such biological shortcuts are presumably
essential to bring into proximity the pointers x . Each of the n MDSs, denoted
primarily by
n , is flanked by the pointers x i 1 ,i and x i,i + 1 . Each
pointer points to the MDS that should precede or follow
α i ,1
i
α i in the final sequence.
The only exceptions are
α 1 , which is preceded by
τ s , and
α n , which is followed
by
τ e in the input string or micronuclear molecule. Note that, although present
generally once in the final macronuclear copy, each x i,i + 1 occurs at least twice
in the micronuclear copy: once after
α i and once before
α i + 1 .
k does not oc-
cur in the final sequence. Thus, since unscrambling leaves one copy of each
x i,i + 1 between
k an internal sequence that is eliminated;
We denote by
k x i,i + 1 or
x i 1 ,i k , depending on which pointer x i,i + 1 is eliminated. Similarly, an MDS is
technically either
α i and
α i + 1 , an IES is nondeterministically either
α i x i + 1 or x i 1 ,i α i . For this model, either choice is equivalent.
The following example (from Landweber and Kari [8]) models unscrambling
of a micronuclear gene that contains MDSs in the scrambled order 2-4-1-3 using
only the operation of linear/circular recombination:
{ ux 12 α 2 x 23 1 x 34 α 4 τ e 2 τ s α 1 x 12 3 x 23 α 3 x 34 v }⇒
{ ux 12 3 x 23 α 3 x 34 v,
•α 2 x 23 1 x 34 α 4 τ e 2 τ s α 1 x 12 }
={ ux 12 3 x 23 α 3 x 34 v,
1 x 34 α 4 τ e 2 τ s α 1 x 12 α 2 x 23 }⇒
{ ux 12 3 x 23 1 x 34 α 4 τ e 2 τ s α 1 x 12 α 2 x 23 α 3 x 34 v }⇒
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