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A more recent version of this article appeared on January 1, 2004
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Submitted on June 17, 2003
Revised on September 15, 2003
Accepted on September 22, 2003
1 Department of Cell Biology and Anatomy, College of Medicine, University of Arizona, Tucson, Arizona 85724, USA
2 Department of Biochemistry, University of Texas Southwestern Medical Center, Dallas, Texas 75390, USA
3 Department of Cell Biology, University of Virginia Health System, Charlottesville, Virginia 22908, USA
* Corresponding author. E-mail address: jeanw{at}email.arizona.edu.
In the developing nervous system, controlled neurite extension and
branching are critical for the establishment of connections between
neurons and their targets. Although much is known about the regulation
of axonal development, many of the molecular events that regulate
axonal extension remain unknown. ARNO and ARF6 have important roles in
the regulation of the cytoskeleton as well as membrane trafficking. To
investigate the role of these molecules in axonogenesis, we expressed
ARNO and ARF6 in cultured rat hippocampal neurons. Expression of
catalytically inactive ARNO or dominant negative ARF6 resulted in
enhanced axonal extension and branching and this effect was abrogated
by coexpression of constitutively active ARF6. We sought to identify
the downstream effectors of ARF6 during neurite extension by
coexpressing PI(4)P 5-Kinase 
with catalytically inactive ARNO and
dominant negative ARF6. We found that PI(4)P 5-Kinase plays a role
in neurite extension and branching downstream of ARF6. Also, expression
of inactive ARNO/ARF6 depleted the actin binding protein Mena from the
growth cone leading edge, indicating that these effects on axonogenesis
may be mediated by changes in cytoskeletal dynamics. These results
suggest that ARNO and ARF6, through PI(4)P 5-Kinase , regulate
axonal elongation and branching during neuronal development.
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