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Sensing the Environment: Regulation of Local and Global Homeostasis by the Skin's Neuroendocrine System PDF

128 Pages·2012·1.86 MB·English
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Reviewsandcriticalarticlescoveringtheentirefieldofnormalanatomy(cytology, histology, cyto- and histochemistry, electron microscopy, macroscopy, experi- mentalmorphologyandembryologyandcomparativeanatomy)arepublishedin AdvancesinAnatomy,EmbryologyandCellBiology.Papersdealingwithanthro- pology and clinical morphology that aim to encourage cooperation between anatomyand related disciplineswillalsobeaccepted. Papers are normally com- missioned. Originalpapers and communications may be submitted and will be consideredforpublicationprovidedtheymeettherequirementsofareviewarticle andthusfitintothescopeof“Advances”.Englishlanguageispreferred. It is a fundamental condition that submitted manuscripts have not been and willnotsimultaneouslybesubmittedorpublishedelsewhere.Withtheacceptance of a manuscript for publication, the publisher acquires full and exclusive copy- rightforalllanguagesandcountries. Twenty-fivecopiesofeachpaperaresuppliedfreeofcharge. Manuscriptsshouldbeaddressedto Co-ordinatingEditor Prof.Dr.H.-W.KORF,ZentrumderMorphologie,Universita¨tFrankfurt,Theodor-SternKai7, 60595Frankfurt/Main,Germany e-mail:[email protected] Editors Prof.Dr.F.BECK,HowardFloreyInstitute,UniversityofMelbourne,Parkville,3000Melbourne,Victoria,Australia e-mail:[email protected] Prof.Dr.F.CLASCA´,DepartmentofAnatomy,HistologyandNeurobiology UniversidadAuto´nomadeMadrid,Ave.ArzobispoMorcillos/n,28029Madrid,Spain e-mail:[email protected] Prof.Dr.D.E.HAINES,Ph.D.,DepartmentofAnatomy,TheUniversityofMississippiMed.Ctr., 2500NorthStateStreet,Jackson,MS39216–4505,USA e-mail:[email protected] Prof.Dr.N.HIROKAWA,DepartmentofCellBiologyandAnatomy,UniversityofTokyo, Hongo7–3–1,113-0033Tokyo,Japan e-mail:[email protected] Dr.Z.KMIEC,DepartmentofHistologyandImmunology,MedicalUniversityofGdansk, Debinki1,80-211Gdansk,Poland e-mail:[email protected] Prof.Dr.R.PUTZ,AnatomischeAnstaltderUniversita¨tMu¨nchen, LehrstuhlAnatomieI,Pettenkoferstr.11,80336Mu¨nchen,Germany e-mail:[email protected] Prof.Dr.J.-P.TIMMERMANS,DepartmentofVeterinarySciences,UniversityofAntwerpen, Groenenborgerlaan171,2020Antwerpen,Belgium e-mail:[email protected] 212 Advances in Anatomy, Embryology and Cell Biology Co-ordinatingEditor H.-W.Korf,Frankfurt Editors F.F.Beck(cid:129)F.Clasca´ (cid:129)D.E.Haines(cid:129)N.Hirokawa Z.Kmiec(cid:129)R.Putz(cid:129)J.-P.Timmermans Forfurthervolumes: http://www.Springer.com/series/102 Andrzej T. Slominski, Michal A. Zmijewski, Cezary Skobowiat, Blazej Zbytek, Radomir M. Slominski, Jeffery D. Steketee Sensing the Environment: Regulation of Local and Global Homeostasis by the Skin’s Neuroendocrine System With 23 figures AndrzejT.Slominski BlazejZbytek UniversityofTennessee UniversityofTennesseeHealth HealthScienceCenter ScienceCenter DepartmentofPathologyandLaboratory DepartmentofPathologyandLaboratory Medicine,andDepartmentofMedicine Medicine Memphis,TN,USA Memphis,TN,USA [email protected] RadomirM.Slominski MichalA.Zmijewski UniversityofTennesseeHealth MedicalUniversityofGdansk ScienceCenter DepartmentofHistology Memphis,TN,USA Gdansk,Poland and CezarySkobowiat JagiellonianUniversityMedicalCollege UniversityofTennessee Krakow,Poland HealthSciencesCenter DepartmentofPathologyandLaboratory JefferyD.Steketee Medicine Memphis,TN,USA UniversityofTennesseeHealth ScienceCenter DepartmentofPharmacology Memphis,TN,USA ISSN0301-5556 ISBN:978-3-642-19682-9 ISBN:978-3-642-19683-6(eBook) DOI:10.1007/978-3-642-19683-6 SpringerHeidelbergNewYorkDordrechtLondon LibraryofCongressControlNumber:2012939853 #Springer-VerlagBerlinHeidelberg2012 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartofthe materialisconcerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,recitation, broadcasting,reproductiononmicrofilmsorinanyotherphysicalway,andtransmissionorinformation storageandretrieval,electronicadaptation,computersoftware,orbysimilarordissimilarmethodology nowknownorhereafterdeveloped.Exemptedfromthislegalreservationarebriefexcerptsinconnection withreviewsorscholarlyanalysisormaterialsuppliedspecificallyforthepurposeofbeingenteredand executed on a computer system, for exclusive use by the purchaser of the work. Duplication of this publicationorpartsthereofispermittedonlyundertheprovisionsoftheCopyrightLawofthePublisher’s location, in its current version, and permission for use must always be obtained from Springer. PermissionsforusemaybeobtainedthroughRightsLinkattheCopyrightClearanceCenter.Violations areliabletoprosecutionundertherespectiveCopyrightLaw. Theuseofgeneraldescriptivenames,registerednames,trademarks,servicemarks,etc.inthispublication doesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexemptfromtherelevant protectivelawsandregulationsandthereforefreeforgeneraluse. Whiletheadviceandinformationinthisbookarebelievedtobetrueandaccurateatthedateofpublication, neithertheauthorsnortheeditorsnorthepublishercanacceptanylegalresponsibilityforanyerrorsor omissionsthatmaybemade.Thepublishermakesnowarranty,expressorimplied,withrespecttothe materialcontainedherein. Printedonacid-freepaper SpringerispartofSpringerScience+BusinessMedia(www.springer.com) Abstract Skin, the body’s largest organ, is strategically located at the interface with the externalenvironmentwhereitdetects,integrates,andrespondstoadiverserange ofstressorsincludingsolarradiation.Ithasalreadybeenestablishedthattheskin is an important peripheral neuro-endocrine-immune organ that is tightly net- worked to central regulatory systems. These capabilities contribute to the main- tenance of peripheral homeostasis. Specifically, epidermal and dermal cells produce and respond to classical stress neurotransmitters, neuropeptides, and hormones. Such production is stimulated by ultraviolet radiation (UVR), biologicalfactors(infectiousandnoninfectious),andotherphysicalandchemical agents. Examples of local biologically active products are cytokines, biogenic amines (catecholamines, histamine, serotonin, and N-acetyl-serotonin), melato- nin, acetylocholine, neuropeptides including pituitary (proopiomelanocortin- derived ACTH, b-endorphin or MSH peptides, thyroid-stimulating hormone) and hypothalamic (corticotropin-releasing factor and related urocortins, thyroid-releasing hormone) hormones as well as enkephalins and dynorphins, thyroid hormones, steroids (glucocorticoids, mineralocorticoids, sex hormones, 7-d steroids), secosteroids, opioids, and endocannabinoids. The production of thesemoleculesishierarchical,organizedalongthealgorithmsofclassicalneuro- endocrine axes such as hypothalamic–pituitary–adrenal axis (HPA), hypotha- lamic–thyroid axis (HPT), serotoninergic, melatoninergic, catecholaminergic, cholinergic,steroid/secosteroidogenic,opioid,andendocannbinoidsystems.Dys- regulationoftheseaxesorofcommunicationbetweenthemmayleadtoskinand/ orsystemicdiseases.Theselocal neuroendocrinenetworksarealsoaddressedat restrictingmaximallytheeffectofnoxiousenvironmentalagentstopreservelocal andconsequentlyglobalhomeostasis.Moreover,theskin-derivedfactors/systems can also activate cutaneous nerve endings to alert the brain on changes in the epidermalordermalenvironments,oralternativelytoactivateothercoordinating centers by direct (spinal cord) neurotransmission without brain involvement. Furthermore, rapidandreciprocalcommunicationsbetweenepidermalandder- mal and adnexal compartmentsare also mediated by neurotransmission includ- ing antidromic modes of conduction. In conclusion, skin cells and skin as an organcoordinateand/orregulatenotonlyperipheralbutalsoglobalhomeostasis. v . Acknowledgments The projects described were supported by grants R01AR052190, R01AR047079, and 1R01AR056666-01A2 from the NIH/NAIMS and grants IBN-9405242, IBN- 9604364, and IOS-0918934 from the NSF to ATS and grants from the Polish Ministry of Science and Higher Education N405 623238 and N402 662840 to MAZ. The chapter on melatonin is dedicated to Dr. Aaron B. Lerner who has discoveredanddefinedthestructureofmelatonin.ThechapteronCRFandCRF receptors is dedicated to Dr. Wylie Vale who codiscovered the CRF. We are grateful to Professor Zbigniew Kmiec for his critical comments, intellectual input,carefuleditorialwork,andtimecommittedtothismanuscript. vii . Contents 1 Introduction .............................................................................. 1 1.1 GeneralOverview .................................................................... 1 1.2 AnOverviewofHistologyandAnatomy .......................................... 3 1.3 AnOverviewofSkinInnervation .................................................. 5 2 BiogenicAminesintheSkin ............................................................ 7 2.1 AnOverview ......................................................................... 7 2.2 Catecholamines ...................................................................... 7 2.2.1 ProductionandMetabolism ................................................. 7 2.2.2 BioregulatoryRoleofCatecholaminesintheSkin ....................... 10 2.3 Histamine .......................................................................... 14 2.3.1 ProductionandMetabolismofHistamine ................................ 14 2.3.2 BioregulatoryRoleofHistamineintheSkin ............................. 15 2.3.3 Conclusions ................................................................. 16 2.4 SerotoninergicSystem ............................................................. 17 2.4.1 ProductionandMetabolismofSerotonin ................................ 17 2.4.2 BioregulatoryRoleofSerotoninintheSkin .............................. 22 2.4.3 SerotoninReceptorsonSensoryNerves .................................. 24 2.4.4 ReceptionofUltravioletLight ............................................. 25 2.4.5 Conclusions ................................................................. 26 3 MelatoninergicSystemintheSkin ................................................... 27 3.1 MelatoninProduction ............................................................. 27 3.2 MelatoninDegradation ............................................................ 29 3.3 BiologicalActivityofMelatonin .................................................. 31 3.4 MelatoninReceptorsintheSkin ................................................. 34 3.5 MelatoninProtectsAgainstSkinDamage ....................................... 35 3.6 Conclusions ........................................................................ 36 4 CutaneousCholinergicSystem ........................................................ 37 4.1 AnOverview ....................................................................... 37 4.2 Conclusions ........................................................................ 38 5 CorticotropinSignalingSystemintheSkin ......................................... 41 5.1 CRFandUrocortins ............................................................... 41 5.2 ExpressionandFunctionsofCRFandUrocortinsintheSkin ................. 41 5.3 SplicingofCRFReceptorTranscripts ............................................ 44 ix

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The skin, the body’s largest organ, is strategically located at the interface with the external environment where it detects, integrates and responds to a diverse range of stressors, including solar radiation. It has already been established that the skin is an important peripheral neuroendocrine-
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