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      Layer- and Cell Type-Specific Modulation of Excitatory Neuronal Activity in the Neocortex

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          Abstract

          From an anatomical point of view the neocortex is subdivided into up to six layers depending on the cortical area. This subdivision has been described already by Meynert and Brodmann in the late 19/early 20. century and is mainly based on cytoarchitectonic features such as the size and location of the pyramidal cell bodies. Hence, cortical lamination is originally an anatomical concept based on the distribution of excitatory neuron. However, it has become apparent in recent years that apart from the layer-specific differences in morphological features, many functional properties of neurons are also dependent on cortical layer or cell type. Such functional differences include changes in neuronal excitability and synaptic activity by neuromodulatory transmitters. Many of these neuromodulators are released from axonal afferents from subcortical brain regions while others are released intrinsically. In this review we aim to describe layer- and cell-type specific differences in the effects of neuromodulator receptors in excitatory neurons in layers 2–6 of different cortical areas. We will focus on the neuromodulator systems using adenosine, acetylcholine, dopamine, and orexin/hypocretin as examples because these neuromodulator systems show important differences in receptor type and distribution, mode of release and functional mechanisms and effects. We try to summarize how layer- and cell type-specific neuromodulation may affect synaptic signaling in cortical microcircuits.

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          Most cited references149

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          International Union of Basic and Clinical Pharmacology. LXXXI. Nomenclature and classification of adenosine receptors--an update.

          In the 10 years since our previous International Union of Basic and Clinical Pharmacology report on the nomenclature and classification of adenosine receptors, no developments have led to major changes in the recommendations. However, there have been so many other developments that an update is needed. The fact that the structure of one of the adenosine receptors has recently been solved has already led to new ways of in silico screening of ligands. The evidence that adenosine receptors can form homo- and heteromultimers has accumulated, but the functional significance of such complexes remains unclear. The availability of mice with genetic modification of all the adenosine receptors has led to a clarification of the functional roles of adenosine, and to excellent means to study the specificity of drugs. There are also interesting associations between disease and structural variants in one or more of the adenosine receptors. Several new selective agonists and antagonists have become available. They provide improved possibilities for receptor classification. There are also developments hinting at the usefulness of allosteric modulators. Many drugs targeting adenosine receptors are in clinical trials, but the established therapeutic use is still very limited.
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            Heterotrimeric G protein activation by G-protein-coupled receptors.

            Heterotrimeric G proteins have a crucial role as molecular switches in signal transduction pathways mediated by G-protein-coupled receptors. Extracellular stimuli activate these receptors, which then catalyse GTP-GDP exchange on the G protein alpha-subunit. The complex series of interactions and conformational changes that connect agonist binding to G protein activation raise various interesting questions about the structure, biomechanics, kinetics and specificity of signal transduction across the plasma membrane.
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              The neural circuit of orexin (hypocretin): maintaining sleep and wakefulness.

              Sleep and wakefulness are regulated to occur at appropriate times that are in accordance with our internal and external environments. Avoiding danger and finding food, which are life-essential activities that are regulated by emotion, reward and energy balance, require vigilance and therefore, by definition, wakefulness. The orexin (hypocretin) system regulates sleep and wakefulness through interactions with systems that regulate emotion, reward and energy homeostasis.
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                Author and article information

                Contributors
                Journal
                Front Neuroanat
                Front Neuroanat
                Front. Neuroanat.
                Frontiers in Neuroanatomy
                Frontiers Media S.A.
                1662-5129
                30 January 2018
                2018
                : 12
                : 1
                Affiliations
                [1] 1Research Centre Jülich, Institute of Neuroscience and Medicine, INM-10 , Jülich, Germany
                [2] 2Department of Psychiatry, Psychotherapy and Psychosomatics, Medical School, RWTH Aachen University , Aachen, Germany
                [3] 3Jülich-Aachen Research Alliance – Translational Brain Medicine , Jülich, Germany
                Author notes

                Edited by: Kathleen S. Rockland, School of Medicine, Boston University, United States

                Reviewed by: Patricia Gaspar, Institut National de la Santé et de la Recherche Médicale, France; Bertrand Lambolez, Université Pierre et Marie Curie, France

                *Correspondence: Dirk Feldmeyer, d.feldmeyer@ 123456fz-juelich.de
                Article
                10.3389/fnana.2018.00001
                5797542
                29440997
                fa099fc4-3b5e-424b-a31d-3a3df347c80e
                Copyright © 2018 Radnikow and Feldmeyer.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 24 August 2017
                : 04 January 2018
                Page count
                Figures: 8, Tables: 1, Equations: 0, References: 163, Pages: 16, Words: 0
                Categories
                Neuroanatomy
                Review

                Neurosciences
                barrel cortex,cortical layers,neuromodulation,acetylcholine,adenosine,dopamine,orexin
                Neurosciences
                barrel cortex, cortical layers, neuromodulation, acetylcholine, adenosine, dopamine, orexin

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