Genes involved in the formation of the earliest cortical circuits.

Department of Physiology, Anatomy and Genetics, University of Oxford, South Parks Road, Oxford OX1 3QX, UK.
Novartis Foundation symposium 02/2007; 288:212-24; discussion 224-9, 276-81.
Source: PubMed

ABSTRACT Building the brain is like erecting a house of cards. The early connections provide the foundation of the adult structure, and disruption of these may be the source of many developmental flaws. Cerebral cortical developmental disorders (including schizophrenia and autism) and perinatal injuries involve cortical neurons with early connectivity. The major hindrance of progress in understanding the early neural circuits during cortical development and disease has been the lack of reliable markers for specific cell populations. Due to the advance of powerful approaches in gene expression analysis and the utility of models with reporter gene expressions in specific cortical cell types, our knowledge of the early cortical circuits is rapidly increasing. With focus on the sub-plate, layer VI and layer V projection neurons, we shall illustrate the progress made in the understanding of their neurochemical properties, physiological characteristics and their integration into the early intracortical and extracortical circuitry. This field benefited from recent developments in mouse genetics in generating models with subtype specific gene expression patterns, powerful cell dissection and separation methods combined with microarray analysis. The emergence of cortical cell type specific biomarkers will not only help neuropathological diagnosis, but will also eventually reveal the causal relations in the pathogenesis of various cortical developmental disorders.

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    ABSTRACT: Development of the cortex involves transient, dynamic cortical circuits, which get substantially remodelled according to the interplay between the developmental programme and the environment. Subplate constitutes a largely transient cortical neuron population. They are among the earliest generated neurons of our brain and lay the foundation of our developing cerebral cortex. Since most initial cortical input and output are directed through subplate neurons, understanding how the connectivity and functional integration of subplate develops, how the transient subplate circuits co-exist with the more permanent cortical networks and how regional variation across the cortex is programmed are issues critical to a broader understanding of cortical function. After subserving these functions, the majority of these fascinating subplate neurons die and give way to the permanent cortical circuits. The developing brain is particularly vulnerable if these transient circuits are damaged or malfunction. Subplate has been implicated in several brain developmental disorders (childhood epilepsy, schizophrenia, autism and cerebral palsy). The inability to identify these cells by criteria other than location in the brain was the major hindrance of progress in the understanding of the role of this enigmatic cell population in cortical development and disease. In this chapter we review the emerging information on the integration of subplate neurons into the cortical and extracortical circuitry using reporter gene expressing mouse transgenic lines; give a flavour of our current research on novel subtype specific markers of selected groups of subplate neurons. These markers started to have an effect in the monitoring of subplate neuronal populations in pathological animal models. KeywordsCerebral cortical development-Subplate neurons-Layer 5 pyramidal neurons-Thalamocortical connections-Golli-tau-eGFP mouse-Migration defects
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