Analysis of replication fork direction in the region downstream of the
3′ regulatory region by N/A 2D gel electrophoresis. The first
dimension of the agarose gel is run in neutral conditions in which
restriction enzyme-digested DNA molecules enriched in replication
intermediates are separated according to their mass. The second
dimension is run in alkaline conditions to separate parental and
nascent strands. The direction of replication fork movement (indicated
by arrow) is determined by hybridization to probes (solid rectangles)
located at the 5′ or 3′ end of a specific restriction fragment (probes
199M11:34 and 40, 73 and
76, and 84 and 88); or from 5′ and 3′
ends of two adjacent restriction fragments (probes 199M11:53
and 57, 60 and 65, 81 and
82, and 92 and 93); or by using a
single probe located at the ends of two overlapping fragments (probes
199M11:79 and 113). The size of the restriction
fragment is indicated. Note that 199M11:34 and
199M11:117 were previously termed Cα-45 and Cα-100,
respectively. Each segment (indicated by gray lines) was analyzed by
two or more 2D gels with similar findings. The topmost restriction
fragment and the topmost arrow (if there are two) always refer to the
left of the two panels. The horizontal line (a) represents
the parental strands of the replication intermediates, which run at a
single position in the second dimension because they have a single
molecular weight. The diagonal line (b) emanating from the
parental strands represents nascent strands of different sizes, ranging
from very small to as large as the parental strands. If replication
forks move from 3′ to 5′, a probe located at the 3′ end of a fragment
will detect nascent strands of sizes ranging from small to
parental-sized, whereas probes located at the 5′ end will detect
nascent strands only of a relatively large size. If forks move in both
directions through a segment, then nascent strands of multiple sizes
will be detected with probes for both 5′ and 3′ ends of this segment.
Because of limitations in the resolution of N/A 2D gel
electrophoresis, we cannot rule out the possibility that <10% of the
forks progress from the opposite direction in the region upstream of
199M11:76. The spots present on the nascent strand arcs and
the linear molecules on the autoradiogram probed with 199M11:84or :88 are caused by the accumulation of a
single-stranded break in this segment, produced by EcoRI
digestion, which is released during alkaline treatment in the 2nd
dimension. No spots were detected on neutral/neutral (N/N) 2D gels.
Therefore, this single-strand break does not appear to form any
replication barrier. H, HindIII; E, EcoRI; B,
BamHI; X, XbaI.