Triple-stranded DNA Discussion on Structure

clive delmonte clived at ndirect.co.uk
Mon Sep 14 08:56:16 EST 1998


Let me thank Eric Fairfield, Kandala Chary and Bill Ross for their helpful,
initial comments made in response to my request on 11th September for
suggestions as to how the problem of triple strand formation inside fibres
might be resolved.

Kandala suggests that:

"In poly(dI).poly(dC) and poly(dA).poly(dT) sequences dI and dA strands have
a propencity to accept the third strand on the major groove side. And in
poly(dI).poly(dC) and poly(dA).poly(dT) mixtures, before they are drawn into
fibres, the duplexes are in dynamic equilibrium with monomeric strands.
Thus, when one forms fibers, the need to unwind a duplex is not necessary.
There are sufficient monomeric strands which hug to the duplexes, thus
forming stable irreversible triplexes. I hope this will answer your
question."

These are very interesting points but it seems to me that there may be some
difficulties with them.  The dynamic equilibrium existing between single
strands and duplexes could be seen to invite a very similar point about how
exactly a duplex unwinds in solution during denaturation.  Where does the
torque come from ?  I have never seen this question addressed.

The unwinding situation in solution is very similar in biophysical terms to
that found in fibres during triple strand formation when one duplex also has
to "unwind", it seems to me.  Moreover, in solution, the duplex is folded
into 3 dimensions, compared to being linear in fibres, and unwinding might
be even harder than in fibres.

As I recall, the major groove does not exist on one side of a duplex, but
follows a helical path around the duplex axis. If a third strand is to join
the first two by lying in the major groove, it would have to be wound onto
the original duplex, I believe.   The initial fibre X-ray diffraction
patterns were characteristic of those ascribed to duplexes and distinct from
single strand diffraction patterns (from poly(rC) and from poly(rA), for
example).  Therefore single strands would seem to have been absent
initially, but were produced in forming the triplex plus a residual single
strand over a period of some days after the young fibre had been drawn.

Bill suggests that:

"Not knowing much about fiber diffraction, it's easy enough to speculate
that somehow the original duplex was somehow not really a duplex, but
assuming it was, I wonder if you are thinking about bond breaking, maybe
some sort of autocatalytic mechanism."

Again, I am grateful for these very interesting suggestions.  Even if the
initial duplexes were not really duplexes, they would seem to have wound
around each other to form the final triplex (assuming that winding around
was involved in triplex formation !)   I think that here lies actually the
germ of the answer.  The initial duplexes were not duplexes in the sense
that the strands were wound around each other, and the final triplex
structures did not involve any strands winding around each other either.
There was no unwinding and no winding either.

This would then become the explanation of denaturation.  The duplexes are
not composed of two strands wound around each other, but two helical strands
running antiparallel alongside each other.  During denaturation the two
helical strands could then move apart in a linear direction.  In triplex
formation in a fibre, one strand moves in a linear manner from its initial
partner to an adjacent duplex. Probably the helical sugar-phosphate chains
would move hardly at all, but the bases might rotate to reach their new
partners.

I wonder if you recall the paper by Kabata et al. (SCIENCE 262 (1993)
1561-1563) ?  They found that single molecules of RNA polymerase were
sliding along tethered molecules of duplex DNA.  The polymerase did not
describe a helical locus at all.  The DNA was immobilised and could not
rotate and nor did the polymerase rotate either.   If duplex DNA consisted
of two chains wound around each other, how would the polymerase "read" the
message on the alternate half turns of the duplex which lay on the other
side away from the polymerase ?  Would that not require two reading
mechanisms, one each to be used on alternating half turns ?  This would
hardly be conducive to high quality transduction of the message contained in
the base sequence.  Kabata et al. did not address this question.   If the
DNA actually consisted of two antiparallel helical chains running alongside
each other, the RNA polymerase could remain on the same face of the duplex
and read the whole message.

Attached to this note should be a diagram of the structure of duplex B-DNA,
if I understand the scanner software, which I believe can account for all
the experimental results with DNA which are known to me and would be very
glad to consider with you what you think its limitations might be. Each
sugar-phosphate helix would have the same diameter as the width of a
Watson-Crick base pair, about 11 Angstroms, and the same pitch as a
Watson-Crick B form double helix, about 34 Angstroms.

 Aaron Klug told Robert Olby in "The Path to the Double Helix", page 373,
that Rosalind Franklin had been working with her Pattersons on separate
helices of diameter 11 Angstroms, which would have been before she could
have known about Watson-Crick base pairs (whose width is necessarily close
to 11 also in order to make the geometry work).


You may know Max Delbruck (Watson's tutor) wrote to Watson in 1953:

 "These (DNA strands) would have to be untwiddled to separate the
threads....In any event, one must postulate that the DNA opens up in some
manner, both for replication and for doing its business otherwise.  In the
structure you describe this opening up is opposed both by the two hydrogen
bonds per nucleotide, and by the interlocking
 of the helices, and it becomes a very important consideration to find a way
out of this dilemma, or to think of a modification of the structure that
does not involve interlocking."

Delbrück's prescience stands out across 40 years and more.

I have been actively scouring the X-ray and NMR literature to bring together
those many experimental results which would seem to support the suggested
side-by-side model.  If you would wish to follow the experimental results
further along what I find a very absorbing avenue of thought do let me know.



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end




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