PMC:529315 / 18285-29535 JSONTXT

{"target":"https://pubannotation.org/docs/sourcedb/PMC/sourceid/529315","sourcedb":"PMC","sourceid":"529315","source_url":"http://www.ncbi.nlm.nih.gov/pmc/529315","text":"Discussion\nRelatively few studies have implicated NT-3 as a chemoattractant agent for sensory and motor axons. Previously noted chemoattractant action of the embryonic mouse maxillary process on trigeminal ganglion neurons (Lumsden and Davies 1986) is now attributed to NT-3 and brain-derived neurotrophic factor (BDNF) (O'Connor et al. 1999). Recently, Tucker et al. (2001) showed that developing sensory and motor axons in limb slice cultures preferentially grow towards neurotrophin-soaked beads instead of following their normal trajectories. Conversely, beads soaked with neurotrophin function-blocking antibodies led to reduction of sensory and motor axon growth towards the limb. In transgenic mice, which over-express NT-3 under the nestin promoter in the central nervous system, the course of the proprioceptive afferents are altered and directed towards the regions with high levels of ectopic NT-3 expression in the spinal cord (Ringstedt et al. 1997). Ringstedt et al. considered the possibility that NT-3 may play a chemoattractant role during the innervation of ventral horns by proprioceptive afferents. However, earlier findings of normal proprioceptive afferent trajectories in chicken embryos despite injection of function-blocking NT-3 antibody into the spinal cord (Oakley et al. 1995) led them to discount this possibility. In that study, though, assays on the effectiveness of antibody perturbation on NT-3-dependent cell survival showed that effectiveness was significant (approximately 90%) but not complete (Oakley et al. 1995). Ringstedt et al. (1997) argue that while central sensory axons may still navigate properly in the absence of NT-3, ectopic expression of NT-3 disrupts their targeting. Recently, another member of the neurotrophin family, BDNF, has been suggested to act as a chemoattractant for sensory axons innervating ear (Tessarollo et al. 2004). In a gene replacement strategy in which BDNF expression was driven by the NT-3 promoter, vestibular axons rerouted towards ectopic sources of BDNF in the cochlea that normally expressed NT-3 and were not innervated by these axons. Ectopic NT-3 supplied using osmotic pumps and adenovirus-mediated expression induce sensory axon growth during regeneration (Zhang et al. 1998, Bradbury et al. 1999, Oudega et al. 1999, Ramer et al. 2002), and induce axonal plasticity of corticospinal axons in injured adult spinal cord (Zhou et al. 2003), where the sprouting axons from the intact site cross the midline towards the NT-3 source on the lesion side of the spinal cord. Our present results are in agreement with these observations, and provide further evidence that NT-3 acts as a chemoattractant for sensory afferents.\nPreviously, normal motor neuron innervation was rescued in NT-3 KO animals by over-expressing NT-3 selectively in their muscles (Wright et al. 1997). It appears that peripheral NT-3 alone is sufficient for rescuing proprioceptive neurons in NT-3 KO animals and proper axonal pathfinding in the spinal cord. However, it is unclear whether NT-3 is absent from the ventral spinal cords of these animals. The possibility has been raised that motor neurons may retrogradely transport NT-3 in muscle to the spinal cord (Chen and Frank 1999). Assays to determine whether any NT-3 is present in the ventral horns of these mice would be informative.\nRecently, Patel et al. (2003) reported their observations on Bax/NT-3 double KO mice they bred. Their results are significantly different from ours. They see no proprioceptive afferents in the periphery of the double knockouts and note that central proprioceptive afferents terminate in the intermediate spinal cord without extending ventrally. Their observations are based on PV immunostaining and DiI labeling of the peripheral nerves. We noted that PV immunolabeling is diminished in our Bax/NT-3 double KOs. In various manipulations of the neurotrophins it has been noted that molecular markers for proprioceptive axons such as PV or calcitonin gene-related peptide for nerve growth factor–responsive axons are compromised (Ringstedt et al. 1997; Patel et al. 2000), thus PV immunostaining in Bax/NT-3 double KOs cannot reveal the extent of proprioceptive axons in the periphery.\nSeventy-three percent of retrogradely labeled gastrocnemius muscle afferents were reported to be expressing TrkC RNA in the adult rat DRG, although some of these (about 10%) may represent cutaneous innervation (McMahon et al. 1994). Presently, it is not clear whether TrkC protein is also made by cutaneous afferents and if so at what stage in development this receptor is expressed by cutaneous axons. Based on the available evidence showing that all proprioceptive neurons are eliminated in NT-3 or TrkC null mice, we think that TrkC staining in the tibial nerve mostly represents proprioceptive axons in the vicinity of their peripheral target. Altogether, our findings suggest a role for NT-3 in initiation of muscle innervation and spindle differentiation by the proprioceptive axons.\nPatel et al. (2003) examined DiI-labeled central DRG axons in the spinal cord of three E17 and two P0 cases, and report that central proprioceptive axons stop in the intermediate laminae, never entering the ventral cord. We have examined nine Bax/NT-3 double KO cases, and often incomplete DiI labeling gives the impression that there are no axons reaching the ventral spinal cord. We have also seen cases similar to theirs (n = 4), but at higher magnification these axons did not have terminal boutons and were not completely labeled. However, with complete fills (n = 5), it was possible to trace these axons into the ventral midline and across to the contralateral side, and visualize terminal boutons at their tips (see Figure 3C). Previously, Arber et al. (2000) reported that members of the Ets family of transcription factors, Er81 and Pea3, are expressed by DRG neurons as well as motor neurons and their target muscle fibers. They found that in the spinal cord of Er81 KO mice, ventral projections of proprioceptive axons were mostly absent, and very few axons made it to the ventral cord. Patel et al. (2003) note that the phenotype they observed in their Bax/NT-3 double KO mice is quite similar to that of Er81 KO mice. They provide evidence that NT-3 induces Er81 expression in DRG explants in vitro. Patel et al. (2003) report that Er81 mRNA expression is diminished (but not abolished) in both NT-3 KO and Bax/NT-3 double KO mice, while their immunohistochemistry shows much less protein expression in the double KO mice. It is highly possible that a small but considerable number of DRG cells express the transcription factor Er81 and that their axons grow beyond the intermediate levels of the spinal cord in Bax/NT-3 double KO mice. While the phenotype of Er81 KO mice is quite dramatic, and most proprioceptive axons stop within the intermediate spinal cord, it is important to note that a few axons still find their way to the ventral spinal cord and target properly to the motor neurons (Arber et al. 2000).\nPatel et al. (2003) also present observations from islet2DTA mice, which lack a significant portion of the motor neurons in the ventral cord. In these mice PV immunostaining shows axons in the ventral horns. Patel et al. argue that since motor neurons are absent in these mice, NT-3 secreted by them could not be a signal for proprioceptive axons to enter the lateral motor columns. However, there is no evidence showing that NT-3 mRNA or protein expressed in the ventral spinal cord is exclusively from motor neurons, and there are no available data indicating that in islet2DTA mice, NT-3 expression in the ventral spinal cord is abolished (Yang et al. 2001; Pun et al. 2002). Studies in embryonic mice reported NT-3 mRNA in the ventral horns of the spinal cord, but it is not definitive that both mRNA and protein are expressed solely by motor neurons. In the adult spinal cord, while motor neurons express high levels of NT-3, other cells, including glia, also express it (Zhou and Rush 1994; Dreyfus et al. 1999; Buck et al. 2000). Our present results, along with those from transgenic mice with NT-3 over-expression in ectopic regions of the spinal cord (Ringstedt et al. 1997), argue for a role of NT-3 in chemoattractant axon guidance of proprioceptive axons in the spinal cord.\nFinally, in culture assays, we see a strong chemoattraction of DRG neurons to localized sources of NT-3. This response is seen in WT, Bax null and in p75 null DRG explants, and in the absence of any other neurotrophins or target-derived axon guidance molecules. Furthermore, in vitro, sensory axon response to NT-3 does not appear to be dose-dependent (Ringstedt et al. 1997; Tucker et al. 2001). NT-3 is capable of attracting axons along distances of up to 1 mm in collagen gel matrix, covering the physiological range it needs to attract axons during development. Previous studies with exogenous or local applications of NT-3 to developing primary sensory axons have indicated that this neurotrophin can attract and induce axonal branching (Ulupınar et al. 2000; Özdinler and Erzurumlu 2001, Özdinler et al. 2004).\nAlong the monosynaptic stretch reflex pathway, only Wnt-3 has been implicated as an axon arborization factor in the spinal cord (Krylova et al. 2002). Another molecule, Slit2, expressed in the midline and by motor neurons (Wang et al. 1999) is capable of inducing axonal branching (Nguyen Ba-Charvet et al. 2001; Özdinler and Erzurumlu 2002). DRG neurons express Robo receptors, which bind to Slits, and proprioceptive axons are therefore capable of responding to Slit signals (Wang et al. 1999). Slit2 does not cause repulsion of NT-3-responsive DRG axons in vitro (Nguyen Ba-Charvet et al. 2001), but causes ectopic branching and arborization of trigeminal axons in the brainstem (Özdinler and Erzurumlu 2002). Thus, Slit2 might also be involved in terminal branching of propioceptive axons in the ventral cord and in the midline branching observed in our double KOs.\nLack of PV expression in Bax/NT-3 KO mice suggests that PV expression might be responsible for proper axon targeting and muscle spindle differentiation. Presently we cannot completely rule out this possibility. However, no defects in axon pathfinding along the monosynaptic reflex arc or in muscle spindle differentiation have been noted in PV KO mice, which develop normally and show no apparent changes in their behavior or physical activity (Schwaller et al. 1999). These observations suggest that axonal targeting defects in Bax/NT-3 double KO mice cannot be simply due to lack of PV expression in proprioceptive cells.\nOur present results suggest that NT-3 acts as a short-range axon guidance cue for proprioceptive axons centrally and peripherally, as they navigate to their targets using other axon guidance cues. In its absence, these axons terminate in inappropriate loci. However, NT-3 may not be the only molecule that plays a role in targeting and terminal branching of sensory axons in the ventral spinal cord. NT-3 most likely acts cooperatively with other axon guidance molecules or by regulating expression of yet to be identified proprioceptive neuron-specific receptors/ligands for numerous axon guidance 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