PMC:1635039 / 23281-29939 JSONTXT

{"target":"https://pubannotation.org/docs/sourcedb/PMC/sourceid/1635039","sourcedb":"PMC","sourceid":"1635039","source_url":"http://www.ncbi.nlm.nih.gov/pmc/1635039","text":"Discussion\nDuring the last few years the Wnt1-Cre transgenic driver line has proven to be a powerful tool for tissue-specific gene deletion in NCCs [12,13,27,28]. Using this approach, several studies have independently shown that the NC-specific deletion of the Tgfbr2 gene leads to a distinct set of calvarial, facial and cardiac defects [8,9,20,29]. Interestingly, these defects appear quite different both in the craniofacial and pharyngeal regions, including the heart, when compared to the corresponding mutants of Alk5, which encodes the TGF-β type I receptor, a prototypical binding partner of TGF-βRII [19] and the present study). While Tgfbr2/Wnt1-Cre mutants as well as mice deficient in Tgf-β2 display the PTA type A4 (truncus arteriosus with interrupted aortic arch [30]), Alk5/Wnt1-Cre mutants reported here demonstrate earlier patterning defects of the PAAs, which is particularly obvious in the 3rd pair of the PAAs. Moreover, the Alk5/Wnt1-Cre mutants display an abnormal patterning of the aortic sac and defective AP septation leading to PTA, reminiscent of type A2 (= truncus artriosus with no main pulmonary artery segment present [30]). However, our results also demonstrate that significant hypoplasia of the aortic sac leads to a severe shortening of the ascending truncal arch, which masks possible defects in derivatives of the 4th PAAs, i.e., interruption of the aortic arch. These observed differences are likely due to substantial apoptosis of Alk5-deficient post-migratory neural crest cells, which is clearly detectable at E10.5, whereas similar intense cell death has not been reported in Tgfbr2/Wnt1-Cre mutants[9].\nOur present results suggest that while TGF-β signaling in cardiac NCCs is predominantly mediated via the ALK5/TGF-βRII receptor complex, ALK5 also mediates signaling of other related ligands, which are either directly or indirectly required for appropriate NCC survival. In fact, it has been shown that, besides TGF-βRII, ALK5 can also form a complex with the Activin type IIB receptor to activate downstream Smads 2/3 [18,31]. Furthermore, a subset of TGF-β-related growth and differentiation factors (GDFs), e.g., GDF8, GDF9, GDF11 and GDF15 could induce these events [17,18,32,33]. Although relevant Gdfs 8, 9 11 and 15 are not expressed in the developing heart, nor do the mice deficient in these Gdfs display developmental cardiac defects, we cannot exclude the possibility that circulating GDFs, perhaps in concert with TGF-βs may contribute to NCC survival during cardiac and pharyngeal morphogenesis.\nWe specifically studied apoptosis at the level of the aortic sac, where the AP-septum forms between the origins of 4th and 6th PAAs. Already at E10.5, we could see intense apoptosis among the postmigratory NCCs in the mesenchyme surrounding the aortic sac at the site where the prospective AP septum forms, i.e., this cell death precedes the AP septal defect seen in mutants. Although some NC-derived cells appeared to be differentiating to smooth muscle cells in the OFT (Fig. 6), we could never detect the AP septum forming in Alk5/Wnt1-Cre mutants between E10.5 and E11.5. These findings suggest that the pool of cells forming the AP septum is severely affected by the cell death. Moreover, it is likely that these cells forming the AP-septum die the before majority of them can differentiate to smooth muscle cells.\nSeveral in vitro studies have suggested an indispensable role for TGF-β-signaling in differentiation of NCCs into smooth muscle cells. Moreover, a recent in vivo study suggested that mice lacking Tgfbr2 in CNCCs display defective NCC differentiation into αSMA-positive cells in the AP septum [9], although this result was later disputed by another study [8]. Our immunohistochemical staining of αSMA in the OFT unequivocally demonstrated that signaling via ALK5 is not required for smooth muscle differentiation in vivo. Moreover, it has been suggested that deletion of Tgfbr2 in NCCs leads to other phenotypic features reminiscent of those seen in the velocardiofacial/DiGeorge syndrome (VCF/DGS) [9] caused by a deletion of the so called DiGeorge critical region (DGCR) on chromosome 22q11 [34,35]. Our present results suggest that although many of the observed phenotypes seen in Alk5/Wnt1-Cre mutants superficially resemble those seen in VCF/DGS, a detailed examination shows that the NC-specific abrogation of Alk5 does not lead to VCF/DGS-like phenotypes. Firstly, while the pharyngeal organ migration fails in Alk5/Wnt1-Cre mutants, perhaps as a result of increased mesenchymal cell death in the pharyngeal region, both the thymus, thyroid and parathyroid seem to develop relatively normally on the histological level in these mutants. Secondly, the NCC death seen in Alk5 mutants affects a predominantly postmigratory population of NCCs, while genes located in the DGCR, i.e., Tbx1 and CrkL, control NCC survival earlier at E8.5-E10 by regulating proliferation of the secondary heart field (SHF), and endoderm expansion, which in turn provides survival signal for NCCs allowing them to populate the pharyngeal region [36-39].\nNCC ablation in the chick has been shown to lead to PTA and to a failure of addition of myocardium from the secondary heart field [40]. It was suggested that the defective migration of cells from the secondary heart field to the OFT in turn resulted in shortening and inappropriate rotation of the OFT, leading to mal-alignment of the arterial pole with the ventricles [41]. While the detected OFT phenotype in Alk5/Wnt1-Cre mutants shared many similarities with that seen in the chick NC ablation models, e.g., PTA and the hypoplastic aortic sac, our current results suggest that the secondary heart field is not severely affected in Alk5 mutants (data not shown). Since we could not detect appropriate rotation of the OFT in neural crest-specific mutants of Alk5 or Alk2, it appears that cells derived from the NC, as well as those from the SHF, are mutually required for proper OFT rotation in mice. However, it appears that these two TGF-β/BMP type I receptors contribute to OFT rotation through different mechanisms. In Alk5 mutants there is very little, if any, detectable septal mesenchyme present, and thus it could be argued that in these mutants OFT rotation fails due to a lack of penetration of septal prongs into the cushion mesenchyme. In contrast, in Alk2/Wnt1-Cre mutants a sizeable septal mesenchyme could be seen, still without any obvious looping of the aortic sac and truncal OFT, suggesting that the mere presence of the septal mesenchyme, without correct smooth muscle cell differentiation, is not sufficient for OFT 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