Principles of Early Vertebrate Forebrain Formation
The formation of the vertebrate central nervous system begins at the onset of gastrulation with the specification of the neuroectoderm or neural plate. This flat sheet of neuroepithelial cells is further patterned along its main axes as it undergoes a com
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Principles of Early Vertebrate Forebrain Formation Florencia Cavodeassi, Tania Moreno-Mármol, María Hernandez-Bejarano and Paola Bovolenta
Abstract The formation of the vertebrate central nervous system begins at the onset of gastrulation with the specification of the neuroectoderm or neural plate. This flat sheet of neuroepithelial cells is further patterned along its main axes as it undergoes a complex morphogenetic reorganisation to give rise to the priomordia of the brain and the spinal cord. In this chapter, we provide a basic overview of the regulatory networks that couple patterning and morphogenesis of the forebrain primordium, which arises from the most anterior part of the neural plate and comprises the telencephalic, retinal, hypothalamic and diencephalic fields. We will describe that, as it occurs in other regions of the developing embryo, morphogenesis and specification of the forebrain primordium is coordinated by a constantly evolving combination of a reduced number of signalling pathways and transcription factors, which together form highly interconnected gene regulatory networks. We will also discuss the still fragmentary information showing that the expression levels of the components of these networks is fine-tuned by different species of non-translated RNAs, which further contribute to originate forebrain complexity from a limited number of key genes. Keywords Transcription factors signalling Forebrain Eye Patterning microRNA
Transcriptional networks Morphogens Cell Retina Telencephalon Cell cohesion
F. Cavodeassi T. Moreno-Mármol M. Hernandez-Bejarano P. Bovolenta (&) Centro de Biología Molecular “Severo Ochoa”, CSIC-UAM, c/Nicolás Cabrera, 1, 28049 Madrid, Spain e-mail: [email protected] F. Cavodeassi T. Moreno-Mármol P. Bovolenta CIBER de Enfermedades Raras (CIBERER), Madrid, Spain © Springer International Publishing Switzerland 2016 J. Castelli-Gair Hombría and P. Bovolenta (eds.), Organogenetic Gene Networks, DOI 10.1007/978-3-319-42767-6_11
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Introduction
The vertebrate central nervous system (CNS) derives from the neuroectoderm, which acquires neural character at the onset of gastrulation. Over subsequent stages of development, the neuroectoderm (or neural plate) undergoes complex morphogenetic reorganisation and progressive patterning to give rise to the brain and the spinal cord. After acquiring neural character, the neuroepithelium is roughly regionalised along the antero-posterior (AP) axis (reviewed in Cavodeassi and Houart 2012; Kiecker and Lumsden 2012). This initial patterning leads to a domain-restricted expression of a number of transcription factors (TFs), at the interface of which, signalling centres are established. These centres influence cell fate of the surrounding tissues and are thus known as secondary organisers in analogy to the primary organizer that influences gastrulation events (Kiecker and Lumsden 2012; Vieira et al. 2010). The factors emanating from the secondary organizers activate additi
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