PMC:1084331 / 1222-5564 JSONTXT

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But whether the orderly expression and activity of transcriptional programs at much later developmental stages, well after cell cycle exit, is an essential step in the progression of neuronal differentiation and circuit assembly has yet to be resolved.\nThe differentiation of sensory neurons of dorsal root ganglia (DRG) has been studied extensively with respect to inductive events that specify neuronal fate [2,3], as well as the involvement of late target-derived neurotrophic factors in the control of neuronal survival [4]. Recent evidence has begun to emerge that target-derived factors are also involved in regulating later aspects of neuronal differentiation [5,6,7]. In particular, genetic experiments have addressed the survival-independent role of neurotrophic factors during development by exploiting strains of mice defective both in neurotrophin signaling and in the function of the proapoptotic gene Bax [8,9]. These studies, for example, have revealed that neurotrophin signaling controls the acquisition of peptidergic traits in nociceptive DRG neurons and the control of target innervation [8,9].\nThe onset of some transcriptional programs in neurons, however, has also been shown to occur long after neurons exit the cell cycle. An emerging principle from work in Drosophila and vertebrates is that target-derived factors play a crucial role in the induction of these transcriptional programs [10]. In Drosophila, retrograde BMP signals from the target region control the terminal differentiation of a subpopulation of peptidergic neurons expressing Apterous and Squeeze [11,12]. In vertebrates, peripheral neurotrophic signals have been shown to direct the onset of expression of the ETS transcription factors Er81 and Pea3 in DRG sensory neurons and motor neuron pools several days after these neurons have become post-mitotic [9,13,14,15,16]. Moreover, the induction of Er81 expression in proprioceptive afferents is known to be mediated by peripheral neurotrophin 3 (NT-3) [9]. These two ETS proteins control late aspects of spinal monosynaptic circuit assembly, with Er81 directing proprioceptive sensory neuron differentiation and Pea3 directing motor neuron pool differentiation, respectively [14,15]. In particular, in the absence of Er81, achieved by mutation in the gene or by deprivation of peripheral neurotrophin signaling, group Ia proprioceptive afferents fail to invade the ventral spinal cord and to make effective synaptic connections with motor neurons [9,14].\nThe involvement of target-derived signals in induction of ETS transcription factor expression raises the question of the necessity for the observed delay in the onset of ETS signaling for neuronal maturation. Would precocious expression of ETS proteins in post-mitotic neurons also direct the appropriate sensory neuron developmental programs? In this study, we have used mouse genetics to test this general idea, by investigating whether the precise timing of onset of ETS transcription factor signaling is essential for normal sensory neuron development. We have assessed the biological effects of inducing ETS signaling either at the correct developmental time, or precociously. We find that within proprioceptive sensory neurons, the late onset of ETS signaling is essential for the establishment of normal sensory afferent projections in the spinal cord. Precocious initiation of ETS signaling in post-mitotic DRG neurons leads to abnormal DRG neuron differentiation characterized by neurotrophin-independent neurite outgrowth and inappropriate profiles of gene expression. Our findings reveal that target-triggered inductive signals provide an effective means of ensuring the late onset of expression of transcription factors, and thus an orderly temporal transcriptional sequence that is crucial for neuronal maturation and circuit 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