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The Reeler Mouse as a Model of Brain Development

Only five years ago, nobody in his right mind would have consid- ered publishing a book on reeler as a model for brain develop- ment. Although this interesting mutation has been with us for half a century, it is fair to say that, in spite of a wave of enthusiasm in the late sixties and early seventies, generated primarily by Sidman, Caviness and colleagues, studies of reeler mice fell pro- gressively out of fashion during the next two decades. All that changed almost overnight when the cloning of the reeler gene, dubbed reelin, was reported in Tom Curran's laboratory in 1995. The fact that the same gene was identified at the same time independently by two other groups using positional cloning sug- gested strongly that reelin was the right candidate. Although the key experiments of transgenic rescue have not been made (and perhaps will never be), the equation "reeler is reelin" has been established beyond reasonable doubt, as alterations of the reelin gene and/or its expression have been found in at least five alleles of reeler and in the mutation Shaking Rat Kawasaki (SRK), an ortholog of reeler.
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Table of Contents

1 Brain Development in Normal and reeler Mice: The Phenotype.- 1.1 Some Introductory Background.- 1.2 The Early Development of the Normal Cerebral Cortex.- 1.2.1 Orientation of Mitoses in the Ventricular Zone.- 1.2.2 The Preplate.- 1.2.3 The Early Marginal Zone.- 1.2.4 The Appearance of the Cortical Plate.- 1.3 Early Cortical Histogenesis in reeler Mutant Mice.- 1.4 The reeler Cerebellum.- 1.5 The reeler Inferior Olivary Complex.- 1.6 The Facial Nerve Nucleus and Other Targets.- 1.6.1 Other Targets of the reeler Gene.- 1.7 ECM Components and Cortical Development.- 1.7.1 ECM and Fiber Growth.- 1.7.2 Integrins.- 2 Genetics of reeler and Genomics of reelin.- 2.1 Genetics and the Various Alleles of reeler.- 2.1.1 Rat reeler-like Mutations.- 2.1.2 Other Mouse Mutations with a reeler-like Phenotype.- 2.2 Mapping of reeler and Cloning of reelin.- 2.2.1 Mapping.- 2.2.2 The reelin cDNA.- 2.3 Genomic Organization of the reelin Gene.- 2.3.1 Alternative Splicing of the reelin Gene.- Alternative Use of a Six Base Pair Microexon.- Alternative Polyadenylation.- 2.3.2 The Promoter Region.- 2.3.3 Clues to the Possible Origin of the reelin Repeats?.- 3 Topography and Cellular Localization of reelin mRNA and Protein Expression During Brain Development.- 3.1 reelin mRNA Expression During Development.- 3.2 Study of Reelin Protein Expression Using Antibodies to Reelin.- 3.2.1 Antireelin Antibodies.- 3.2.2 Reelin Protein Expression During Mouse Brain and Human Cortical Development.- 3.2.3 Studies of Reelin Function Using Antireelin Antibodies.- 3.2.4 Is the Reelin Protein Processed in the Embryonic Mouse Brain?.- 3.3 Some Questions Raised by the Studies of Reelin Expression.- 3.3.1 Cajal-Retzius Cells and Other Reelin-positive Cells in the Developing Cortical Marginal Zone.- 3.3.2 Does Reelin Act on Postmigratory Neurons, Radial Glia or Both?.- 3.3.3 Reelin Expression is Poorly Correlated With the reeler Phenotype.- 3.3.4 Reelin as a Repulsive, Extracellular Matrix-Expanding Molecule?.- 3.3.5 Reelin and Axonal Growth.- 4 The reeler Mutation and Brain Evolution.- 4.1 The Evolution of Brain Development: A New Theme?.- 4.2 Comparative Data on Cell Migration, Maturation, Synaptogenesis And Neurogenesis.- 4.3 Comparison of Cortical Plate Development in Emys and Lacerta.- 4.3.1 Biological Mechanisms Involved in the Histogenesis of the Cortical Plate.- 4.3.2 Evolutionary Considerations.- 4.4 Reelin and a Model of Cortical Evolution.- 5 A Model of Cortical Development Inspired by reeler: Facts and Hypotheses.- 5.1 Early Hypotheses on the Actions of the reeler Gene on the Developing Brain.- 5.1.1 Reeler and Cell Interaction/Adhesion.- 5.1.2 The Action of the reeler Gene is Intrinsic to the Neuroepithelium.- 5.1.3 Reeler Phenotype and Radial Glial Fibers.- 5.2 Mouse Disabledl and the Scrambler/yotari Mutations.- 5.3 Cyclin-Dependent Protein Kinase 5 and its Activator p35: Definition of a New Step in Mammalian Cortical Development.- 5.4 Reeler-Type Malformations and Human Neuropathology.- References.

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