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Volume 16, Number 10,
Issue of May 15, 1996
pp. 3274-3286
Copyright ©1996 Society for Neuroscience
Experience-Dependent Plasticity of Binocular Responses in the
Primary Visual Cortex of the Mouse
Received Nov. 8, 1995; revised Feb. 7, 1996; accepted Feb. 19, 1996.
Joshua A. Gordon and
Michael P. Stryker
Neuroscience Graduate Program, and W. M. Keck Foundation Center for
Integrative Neuroscience, Department of Physiology, University of
California, San Francisco, California 94143-0444
An activity-dependent form of synaptic plasticity underlies the
fine tuning of connections in the developing primary visual cortex of
mammals such as the cat and monkey. Studies of the effects of
manipulations of visual experience during a critical period have
demonstrated that a correlation-based competitive process governs this
plasticity. The cellular mechanisms underlying this competition,
however, are poorly understood. Transgenic and gene-targeting
technologies have led to the development of a new category of reagents
that have the potential to help answer questions of cellular mechanism,
provided that the questions can be studied in a mouse model. The
current study attempts to characterize a developmental plasticity in
the mouse primary visual cortex and to demonstrate its relevance to
that found in higher mammals. We found that 4 d of monocular lid suture
at postnatal day 28 (P28) induced a maximal loss of responsiveness of
cortical neurons to the deprived eye. These ocular dominance shifts
occurred during a well-defined critical period, between P19 and P32.
Furthermore, binocular deprivation during this critical period did not
decrease visual cortical responses, and alternating monocular
deprivation resulted in a decrease in the number of binocularly
responsive neurons. Finally, a laminar analysis demonstrated plasticity
of both geniculocortical and intracortical connections. These results
demonstrate that an activity-dependent, competitive form of synaptic
plasticity that obeys correlation-based rules operates in the
developing primary visual cortex of the mouse.
Key words:
ocular dominance;
neural plasticity;
visual cortex;
development;
mouse;
monocular deprivation;
critical period;
binocular
deprivation
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|
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 |
|

|
 |

|
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|
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|
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