Abstract
The olfactory epithelium (OE) and its associated perireceptor space, i.e., the mucus layer (ML) covering the epithelium, are the most peripheral parts of the vertebrate olfactory system. The olfactory receptor neurons (ORNs), one of the cell types of the OE, are the odorant detectors of the olfactory system. These bipolar neurons extend their apical appendages, which express odorant receptors, into the ML. The binding of odorants to odorant receptors is the initial step of odor processing. The vast majority of research on the peripheral olfactory system has focused on the ORNs and the molecular components of the olfactory transduction cascades. Less attention has been directed to the other cell types of the OE and their physiological functions. For a long time, it was assumed that the olfactory signals detected in the OE are transmitted to the olfactory bulb without preprocessing, but this view turned out to be over-simplistic. It has been shown that the olfactory signals are critically modulated already in the OE. Despite compelling evidence, many descriptions of the olfactory system still ignore the existence of these peripheral modulatory mechanisms. The importance of peripheral modulation of the olfactory signals, the physiological functions of the other epithelial cell types, the extrinsic innervation of the olfactory mucosa, and the perireceptor space are only slowly coming into focus in the olfactory research. Furthermore, several intraepithelial signaling pathways that signal epithelial damage and initiate regenerative processes have recently been discovered. This review provides a concise overview of the current knowledge of peripheral events in the olfactory mucosa and the perireceptor space.
Zusammenfassung
Das olfaktorische Epithel und der zugehörige Perirezeptorraum, d. h. die Schleimschicht, die das Epithel bedeckt, sind die peripheren Teile des olfaktorischen Systems der Vertebraten. Die olfaktorischen Rezeptorneuronen, eine der Zellarten des olfaktorischen Epithels, sind die Geruchsdetektoren des olfaktorischen Systems. Diese bipolaren Neuronen erstrecken ihre apikalen Fortsätze, die Geruchsrezeptoren exprimieren, in die Schleimschicht. Die Bindung von Geruchsstoffen an Geruchsrezeptoren ist der erste Schritt der Geruchsverarbeitung. Der Großteil der Forschung über das periphere olfaktorische System hat sich auf die olfaktorischen Rezeptorneuronen und die molekularen Komponenten der olfaktorischen Transduktionskaskaden konzentriert. Den anderen Zelltypen des olfaktorischen Epithels und ihren physiologischen Funktionen wurde weit weniger Aufmerksamkeit geschenkt. Lange Zeit ging man davon aus, dass olfaktorische Signale, die im olfaktorischen Epithel erkannt werden, ohne Vorverarbeitung an den Bulbus olfactorius weitergeleitet werden, doch diese Ansicht erwies sich als zu vereinfacht. Es hat sich gezeigt, dass olfaktorische Signale bereits im Epithel entscheidend moduliert werden. Trotz überzeugender Beweise vernachlässigen viele Beschreibungen des olfaktorischen Systems immer noch die Existenz dieser peripheren Modulationsmechanismen. Erst sehr langsam rückt die Bedeutung der peripheren Modulation von olfaktorischen Signalen, die physiologischen Funktionen der anderen Zelltypen des olfaktorischen Epithels, die extrinsische Innervation der olfaktorischen Mukosa und der Perirezeptorraum in den Fokus der Riechforschung. Darüber hinaus wurden in den letzten Jahren mehrere intraepitheliale Signalwege entdeckt, die Epithelschäden signalisieren und Regenerationsprozesse einleiten. Dieser Übersichtsartikel gibt einen prägnanten Überblick über den aktuellen Wissensstand zu peripheren Ereignissen in der olfaktorischen Mukosa und dem Perirezeptorraum.
Funding source: Deutsche Forschungsgemeinschaft http://dx.doi.org/10.13039/501100001659
Award Identifier / Grant number: 4113/4-1
About the author
Ivan Manzini is professor of Animal Physiology at the Justus-Liebig-University of Gießen, Germany. The research activities of his group primarily focus on developmental, molecular, cell biological, and behavioral aspects of the olfactory system of the African Clawed Frog, Xenopus laevis. He studied biology at the University of Modena and Reggio Emilia, Italy and received his PhD in Neuroscience from the University of Göttingen in 2003. He, then, conducted a postdoctoral research in the group of Prof. Dr. Dr. Detlev Schild (University of Göttingen, Germany). From 2010 to 2017, he led an independent research group at the DFG Research Center 103—Cluster of Excellence 171—Nanoscale Microscopy and Molecular Physiology of the Brain (CNMPB) at the University of Göttingen, Göttingen, Germany.
Acknowledgment
I would like to thank Dr. Thomas Hassenklöver for reading the manuscript and providing a number of helpful comments.
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Author contributions: The author has accepted responsibility for the entire content of this submitted manuscript and approved submission.
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Research funding: This work was supported by DFG Grant 4113/4-1.
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Conflict of interest statement: The author declares no conflicts of interest regarding this article.
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