PMC:1208873 / 24624-31351 JSONTXT

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Instead, our results reveal that huntingtin is required for normal anterior streak formation and the consequent production of paraxial mesoderm, with a previously unrecognized role for huntingtin in the proper extinction of transiently and/or dynamically expressed genes.\nIndeed, the hallmark of the huntingtin deficient molecular phenotype is the impaired down-regulation of a subset of dynamically expressed genes, after the proper onset of expression. This phenomenon does not reflect a lack of anterior/posterior axis formation, as evidenced by the formation of the AVE anteriorly and the primitive streak posteriorly. Nor can it be simply explained by delayed development, as stage-specific markers, such as Wnt3a and primordial germ cells, which are detectable at E7.0 in wild-type embryos, are induced appropriately. Furthermore, the expression of T and Evx1 in the extraembryonic mesoderm of mutant embryos is not a feature of wild-type embryos, even at earlier stages. This strongly suggests that in huntingtin deficient embryos, the migration of the distal streak derivatives to the extraembryonic mesoderm occurs normally but that the down-regulation of these genes is impaired. This impairment may also explain the failure of huntingtin deficient embryos to properly restrict the expression of Fgf8, Nodal and Gsc. Thus, huntingtin may play a direct role in the transcriptional regulation, or mRNA stability of these genes or it may act indirectly by intersecting with other pathways that regulate the expression of these genes.\nThe requirement for huntingtin in the extraembryonic tissues had prompted us to test whether impaired extraembryonic signals might be responsible for the dysregulation of gene expression within the epiblast that is observed in Hdhex4/5/Hdhex4/5 embryos. Extraembryonic development in Hdhex4/5/Hdhex4/5 embryos is associated with mildly elevated levels expression of Hnf4 in the primitive endoderm and Pem in the lopsided anterior chorion but the expression of other known signals, such as Bmp4 from the extraembryonic ectoderm, and Dkk1 and Lefty1 from the AVE, appear to be normal, although the slight increase in Dkk-1 expression in Hdhex4/5/Hdhex4/5 embryos suggests that further investigation into Wnt signaling is warranted. Moreover, extraembryonic Bmp4 signaling is not impaired in the absence of huntingtin, as the induction of PCGs in mutant embryos is normal, implying proper transport and secretion of the appropriate extraembryonic signals. However, Nodal, Fgf8 and Gsc are expressed ectopically in the visceral endoderm of Hdhex4/5/Hdhex4/5 embryos. Both Nodal and Fgf8, important growth factors required for normal development of the epiblast, are tightly regulated during gastrulation. Therefore, misexpression of either or both of these factors, or of goosecoid, in the visceral endoderm could contribute to the Hdhex4/5/Hdhex4/5 mutant phenotype. In addition, it is possible that other extraembryonic signal(s) that we have not analyzed may also be affected by the lack of huntingtin activity in extraembryonic cells in mutant embryos.\nHuntingtin deficient embryos also fail to form headfolds, and to undergo organogenesis, even after culturing in nutrient rich media. The absence of headfold formation in these embryos does not appear to be a result of a failure to induce neurectoderm or a failure to form the AVE, since mutant embryos express markers such as Otx2, Ddk1, Lefty1 and Hesx1. In addition, since node formation is not required for neural induction [44-46], the failure to form a node in huntingtin deficient embryos is also unlikely to explain the lack of headfolds. The apparent reduction of paraxial mesoderm in Hdhex4/5/Hdhex4/5 embryos could explain the lack of headfolds since paraxial mesoderm is important for the full development of neuroectoderm, and consequently, headfolds. Alternatively, the inability to manifest headfolds could suggest that huntingtin is required at a very early stage for normal CNS development. This conclusion is consistent with the finding that severely reduced levels of huntingtin, from a hypomorphic Hdh allele, lead to abnormal brains later in embryonic development [15].\nThe cardinal features of complete Hdh inactivation that we observe are similar to the phenotypes that stem from the complete inactivation of the Polycomb group gene (Pc-g) Eed (embryonic ectoderm development). Indeed, complete deficiency for either huntingtin or the eed protein leads to abnormal streak development, lack of headfold formation, ectopic T, Evx1 and Nodal expression and disruption of anterior primitive streak mesoderm production [47]. Interestingly, Eed protein is also required for proper trophoblast development and normal maintenance of imprinted X-inactivation and genomic imprinting [47-49], suggesting that these activities warrant investigation in huntingtin deficient embryos.\nThus, our observations provide unexpected starting-points in the search for huntingtin's precise molecular activities, which began with the discovery that this HEAT domain protein hosts the dominant polyglutamine property that is the fundamental basis of HD pathogenesis. In HD patients and in accurate genetic replicas, HD CAG knock-in mice, the dominant mutation specifically affects the major population of neurons in the striatum, without impairing huntingtin's essential activities in embryonic development [50-53]. Indeed, homozygous HD patients make no wild-type huntingtin, and, in the mouse, a single mutant Hdh allele's worth of mutant huntingtin can fully rescue huntingtin deficiency embryonic phenotypes [15,51]. The quest to understand the HD mechanism, therefore, is aimed at delineating the huntingtin activity that may explain the striatal cell specificity of the polyglutamine mutant version of huntingtin. One hypothesis is that huntingtin is normally involved in gene transcription, as proposed for NRSF/REST mediated BDNF expression [54]. Now, our finding that huntingtin can be absolutely necessary for the appropriate regulation of genes with dynamic expression patterns in vivo, provides a compelling reason to elucidate the cellular machinery that is necessary for huntingtin mediated gene 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