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The Potential Role of Rac Signalling and the Planar Cell Polarity Pathway in Wiring of the Enteric Nervous System

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The Potential Role of Rac Signalling and the Planar Cell Polarity Pathway in Wiring of the Enteric Nervous System The potential role of Rac signalling and the Planar Cell Polarity pathway in wiring of the Enteric Nervous System Valentina Sasselli February 2011 Division of Molecular Neurobiology MRC National Institute for Medical Research The Ridgeway Mill Hill, London NW7 1AA Department of Cell and Developmental Biology University College London Thesis submitted to the University College London for the degree of Doctor of Philosophy Declaration of authenticity This work has been completed in the laboratory of Vassilis Pachnis, in the Division of Molecular Neurobiology at the MRC National Institute for Medical Research, London. I, Valentina Sasselli declare that the work presented in this thesis is the result of my own independent investigation. Where information has been derived from other sources, I confirm that this has been indicated in the thesis. The analysis of Rac1 gene expression on Sox10Cre;Rac1flox embryos (section 3.1.1) was performed in collaboration with Dr Silvia Bogni (former post-doc in the laboratory, now at NeuroGlia Unit, San Raffaele Scientific Institute, Italy). I was supported by the Medical Research Council. Valentina Sasselli February, 2011 2 Abstract The functional development of the enteric nervous system (ENS) requires newly generated neurons and their progenitors to migrate to their appropriate sites, extend neurites, guide axons and dendrites to suitable locations and establish synaptic connections with the appropriate targets. Very little is known about the molecular mechanism underlying these processes. Recent studies have suggested a potential role of Rho GTPases as intracellular regulators of several ENS developmental processes. However, the relative participation of specific members of the family in migration, neurogenesis and axonal guidance of enteric progenitors has not been addressed yet. Here, we investigate the in vivo role and genetic interaction of two members of the Rho-GTPase family, Rac1 and Rac3 in enteric neurogenesis. Taking advantage of the Cre/loxP recombination system and a Rac1 conditional inactivation mouse strain (Rac1flox/flox), we generated a Sox10Cre; Rac1flox/flox;R26ReYFP mouse line, where Rac1 gene is specifically ablated in the neural crest population which is also labeled by the expression of Yellow Fluorescent Protein. Secondly, we generated double Rac1;Rac3 mutant animals by crossing the Sox10Cre;Rac1flox;R26ReYFP mouse line to a constitutive Rac3 KO strain (Rac3-/-). In vivo and in vitro studies on Rac-deficient enteric neural crest cells and neurons showed distinctive roles for Rac1 and Rac3 in migration of enteric neural crest cells (ENCCs), in development of enteric neurons and in control of cell polarity within the developing ENS. In addition, we also undertook a candidate gene approach to investigate the involvement of Wnt-signaling genes in enteric axon guidance and circuit formation. We found that two of the core components of the Planar Cell Polarity pathway, the Wnt receptor Frizzled 3 (Fzd3) and the Cadherin EGF LAG seven-pass G-type receptor 3 (Celsr3) are expressed specifically in ENCCs during embryonic development. Here we show, by using a combination of in vivo approaches that in mice deficient in either protein, enteric neurons had characteristic defects in neuronal tract formation and in patterning of individual axonal projections evident from early stages of ENS development. Furthermore, preliminary data show that these specific defects in ENS wiring might be the cause of impaired intestinal function and, therefore, provide the basis for understanding the aetiopathology of several idiopathic enteric neuropathies in humans. 3 Table of contents Title.....................................................................................................................................1 Declaration of authenticity...............................................................................................2 Abstract..............................................................................................................................3 Table of contents ...............................................................................................................4 List of figures.....................................................................................................................8 List of tables.....................................................................................................................10 Abbreviations ..................................................................................................................11 Acknowledgements .........................................................................................................14 CHAPTER 1 Introduction......................................................................... 17 1.1 Enteric nervous system anatomy and function..............................................18 1.1.1 Anatomy and organisation..................................................................................... 18 1.1.2 Cellular components of the ENS: enteric glial cells .............................................. 20 1.1.3 Cellular components of the ENS: enteric neurons................................................. 22 1.1.3.1 Morphology....................................................................................................... 24 1.1.3.2 Electrophysiology ............................................................................................. 24 1.1.3.3 Chemical coding ............................................................................................... 25 1.1.3.4 Functional classes of enteric neurons............................................................... 25 1.1.3.5 Cellular targets ................................................................................................. 27 1.1.4 Physiology of the ENS........................................................................................... 27 1.2 Enteric nervous system development .............................................................29 1.2.1 Pre-enteric neural crest cells.................................................................................. 29 1.2.2 Enteric neural crest cells........................................................................................ 32 1.2.3 Molecular mechanisms controlling the colonisation of the gut by ENCCs........... 35 1.2.3.1 Cell number....................................................................................................... 35 1.2.3.2 Proliferation and survival................................................................................. 36 1.2.3.3 Migration .......................................................................................................... 40 1.2.4 Development of enteric neurons and ENS patterning............................................ 45 1.2.4.1 Neuronal subtype specification......................................................................... 45 1.2.4.2 Morphological development of enteric neurons ............................................... 49 1.2.4.3 Axon guidance................................................................................................... 50 1.2.5 Development of gut motility.................................................................................. 52 4 1.3 Enteric neuropathies........................................................................................54 1.3.1 Congenital enteric neuropathies............................................................................. 55 1.4 Rho GTPases and nervous system development ...........................................59 1.4.1 Rho and Rac GTPases............................................................................................ 59 1.4.2 Rac GTPases and neuronal development............................................................... 60 1.4.2.1 Migration of progenitor cells and neurons ....................................................... 61 1.4.2.2 Neuronal morphology and neuritogenesis........................................................ 62 1.4.2.3 Axon guidance................................................................................................... 63 1.4.3 Rac GTPases in ENS development........................................................................ 65 1.5 Wnt signalling in neural circuit development ...............................................68 1.5.1 Canonical and non-canonical Wnt pathways......................................................... 68 1.5.2 PCP genes and neuronal development................................................................... 70 1.5.2.1 Migration .......................................................................................................... 71 1.5.2.2 Neuronal morphology and neuritogenesis........................................................ 72 1.5.2.3 Axon guidance................................................................................................... 73 1.5.3 Wnt signalling in ENS development ..................................................................... 74 1.6 Aims of the present work.................................................................................76 CHAPTER 2 Materials and Methods ...................................................... 77 2.1 Materials ...........................................................................................................78 2.1.1 Solutions and buffers ............................................................................................
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