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Differentiated neurons balance the need to maintain a stable identity with their flexible responses to dynamic environmental inputs. Here we characterize these opposing influences on gene expression in  Caenorhabditis elegans  olfactory neurons. Using transcriptional reporters that are expressed differentially in two olfactory neurons, AWC ON  and AWC OFF , we identify mutations that affect the long-term maintenance of appropriate chemoreceptor expression. A newly identified gene from this screen, the conserved transcription factor  hmbx-1 , stabilizes AWC gene expression in adult animals through dosage-sensitive interactions with its transcriptional targets. The late action of  hmbx-1  complements the early role of the transcriptional repressor gene  nsy-7 : Both repress expression of multiple AWC OFF  genes in AWC ON  neurons, but they act at different developmental stages. Environmental signals are superimposed onto this stable cell identity through at least two different transcriptional pathways that regulate individual chemoreceptor genes: a cGMP pathway regulated by sensory activity, and a  daf-7  (TGF-β)/ daf-3  (SMAD repressor) pathway regulated by specific components of the density-dependent  C. elegans  dauer pheromone.

The homeodomain protein hmbx-1 maintains asymmetric gene expression in adult C. elegans olfactory neurons