PMC:1208873 / 2123-6184 JSONTXT

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This observation strongly suggests that the striatal cell specificity of the polyglutamine expansion in the context of huntingtin must be determined by some aspect of huntingtin's structure, subcellular location or activities [2].\nHuntingtin is postulated to function as a flexible ~350 kDa HEAT domain scaffold that may facilitate the assembly and possibly the subcellular location of large protein complexes [3-7]. Huntingtin's large number of diverse cytoplasmic and nuclear protein binding partners strongly suggest that huntingtin may participate in a variety of cellular processes that range from trafficking of growth factor complexes to gene transcription (reviewed in [5,8,9]. However, despite the potential importance of huntingtin's normal function to our understanding of how the dominant polyglutamine mutation causes HD pathology, huntingtin's precise molecular and cellular activities have not been defined.\nTherefore, we, and others, set out to discover huntingtin's essential activities by studying the effects of huntingtin deficiency in the mouse. Inactivation of the mouse HD gene (Hdh) has shown that huntingtin is not required for cell viability, as evidenced by the survival of mouse embryonic stem cells and neurons that lack huntingtin [10-12]. However, huntingtin is needed at the level of the organism for proper mammalian embryonic development [10,13,14]. Complete lack of huntingtin results in developmental arrest during gastrulation, while severe reduction of huntingtin levels results in abnormal neurogenesis and perinatal lethality [15].\nAnalysis of huntingtin deficient Hdhex4/5/Hdhex4/5 embryos reveals that homozygous inactivation of the mouse HD gene does not overtly affect development until E7.0. By E7.5, mutant embryos exhibit a shortened primitive streak, reduced size and, by morphology, lack a node and head folds. Mutants are rapidly resorbed by E8.0 [10]. Importantly, the expression of huntingtin only in extraembryonic tissues in chimeras rescues this gastrulation phenotype, suggesting that huntingtin is required only in cells of the extraembryonic lineage and acts in a cell non-autonomous manner at this stage [16].\nExtraembryonic tissues are essential for supplying nutrients and signals that direct anterior/posterior axis formation and patterning in the developing embryo (reviewed in [17]), implicating huntingtin in either or both of these processes. Of these possibilities, the nutritive role has been more extensively investigated. However, huntingtin deficient embryos do not display obvious visceral endoderm defects, with the notable exception of compromised iron transport in later stage mutants, although iron uptake is undisturbed [16] and endocytosis is not impaired in huntingtin deficient embryos or embryonic stem cells [16,18].\nBy the same token, huntingtin shuttles through the nucleus, where it is required for proper nuclear localization of its transcription factor partners, suggesting that huntingtin may play a role in transcription cascades in extraembryonic tissues that pattern the embryo [18]. Therefore, we have examined this hypothesis, by monitoring the expression of genes that determine normal embryonic patterning and morphogenesis in Hdhex4/5/Hdhex4/5 huntingtin deficient embryos. Our results support and refine the hypothesis, indicating that huntingtin is required for proper mesoderm patterning and for normal regional restriction of the expression of a subset of growth and transcription 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