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-related receptor
signaling in the Caenorhabditis elegans
dauer pathway
Department of Molecular Biology, Massachusetts General Hospital,
Department of Genetics, Harvard Medical School,
Boston, Massachusetts 02114 USA
Signals from TGF-
superfamily receptors are transduced to the
nucleus by Smad proteins, which transcriptionally activate target
genes. In Caenorhabditis elegans, defects in a TGF--related pathway cause a reversible developmental arrest and metabolic shift at
the dauer larval stage. Null mutations in daf-3 suppress mutations in genes encoding this TGF- signal, its receptors, and
associated Smad signal transduction proteins. daf-3 encodes a
Smad protein that is most closely related to mammalian DPC4, and is
expressed throughout development in many of the tissues that are
remodeled during dauer development. DAF-4, the type II TGF- receptor
in this pathway, is also expressed in remodeled tissues. These data
suggest that the DAF-7 signal from sensory neurons acts as a
neuroendocrine signal throughout the body to directly regulate
developmental and metabolic shifts in tissues that are remodeled during
dauer formation. A full-length functional DAF-3/GFP
fusion protein is predominantly cytoplasmic, and this localization is
independent of activity of the upstream TGF--related pathway.
However, this fusion protein is associated with chromosomes in mitotic
cells, suggesting that DAF-3 binds DNA directly or indirectly. DAF-3
transgenes also interfere with dauer formation, perhaps attributable to
a dosage effect. A truncated DAF-3/GFP fusion protein
that is predominantly nuclear interferes with dauer formation, implying
a role for DAF-3 in the nucleus. These data suggest that DAF-7 signal
transduction antagonizes or modifies DAF-3 Smad activity in the nucleus
to induce reproductive development; when DAF-7 signals are disabled,
unmodified DAF-3 Smad activity mediates dauer arrest and its associated
metabolic shift. Therefore, daf-3 is unique in that it is
antagonized, rather than activated, by a TGF- pathway.
[Key Words:
diapause; neuroendocrine; TGF-
signaling]
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