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00245_Field_frep126.txt
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1996-12-30
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The final pattern of left-eye,
right-eye alternation in
cortical layer 4C develops
normally even if both eyes are
sewn shut, indicating that the
appropriate wiring can come
about in the absence of
experience. We suppose that
during development, the
incoming fibers from the two
eyes compete in layer 4C in
such a way that if one eye has
the upper hand at any one
place, the eye's advantage, in
terms of numbers of nerve
terminals, tends to increase,
and the losing eye's terminals
correspondingly recede. Any
slight initial imbalance thus
tends to increase progressively
until, at age one month, the
final punched-out stripes
result, with complete
domination everywhere in
layer 4. In the case of eye
closure, the balance is
changed, and at the borders of
the stripes, where normally the
outcome would be a close battle,
the open eye is favored and
wins out, as shown in the
diagram to the left.
This competition model
explains the segregation of
fourth-layer fibers into eye-
dominance columns. At birth
the columns have already
begun to form. Normally at any
given point if one eye
dominates even slightly, it ends
up with a complete monopoly. If
an eye is closed at birth, the
fibers from the open eye still
surviving at any given point in
layer 4 take over completely.
The only regions with
persisting fibers from the
closed eye are those where that
eye had no competition when it
was closed.
We don't know what causes
the initial imbalance during
normal development, but in
this unstable equilibrium
presumably even the slightest
difference would set things off.
Why the pattern that develops
should be one of parallel
stripes, each a half-millimeter
wide, is a matter of speculation.
An idea several people espouse
is that axons from the same eye
attract each other over a short
range but that left-eye and
right-eye axons repel each
other with a force that at short
distances is weaker than the
attracting forces, so that
attraction wins. With
increasing distance, the
attracting force falls off more
rapidly than the repelling
force, so that farther away
repulsion wins. The ranges of
these competing tendencies
determine the size of the
columns. It seems from the
mathematics that to get parallel
stripes as opposed to a
checkerboard or to islands of
left-eye axons in a right-eye
matrix, we need only specify
that the boundaries between
columns should be as short as
possible.
One thus has a way of
explaining the shrinkage and
expansion of columns, by
showing that at the time the
eye was closed, early in life,
competition was, after all,
possible.