An Introduction To Neuroendocrinology

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Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationAn Introduction to NeuroendocrinologySecond EditionHow does the brain regulate sexual behavior, or control our body weight? How do wecope with stress? Addressing these questions and many more besides, this thoroughlyrevised new edition reflects the significant advances that have been made in the study ofneuroendocrinology over the last 20 years.The text examines the importance of the hypothalamus in regulating hormone secretion from the endocrine glands, describing novel sites of hormone release, includingbone, heart, skeletal muscle, and liver. The role of steroid hormone, neurotransmitter andpeptide receptors, and the molecular responses of target tissues, is integrated into thediscussion of the neuroendocrine brain, especially through changes in gene expression.Particular attention is attached to neuropeptides, including their profound influence onbehavior.Complete with new full-color figures throughout, along with review and essay questionsfor each chapter, this is an ideal resource for undergraduate and graduate students ofneuroscience, psychology, biology, and physiology.Michael Wilkinson has 40 years of experience in teaching neuroscience andneuroendocrinology to undergraduate and graduate students as a Professor in theDepartment of Obstetrics and Gynaecology and IWK Health Centre, Dalhousie University,Halifax, Canada. His research laboratory has focused on neurodevelopmental aspectsof female reproduction with a specific interest in the neuroendocrine regulation ofhypothalamic function, including the impact of sex hormones on sleep.Richard E. Brown is a University Research Professor in the Department of Psychologyand Neuroscience at Dalhousie University. He has taught courses on hormones and behavior, measuring behavior, and the neurobiology of learning and memory for more than35 years. His research is on mouse models of Alzheimer’s Disease, Fragile X Syndrome,ADHD, and other neurological disorders. He is currently examining the age-related hormonal changes in transgenic Alzheimer’s mice. in this web service Cambridge University Presswww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationAn Introduction to in this web service Cambridge University Presswww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationNeuroendocrinologySecond EditionMichael WilkinsonProfessor of Obstetrics & GynaecologyProfessor of Physiology & BiophysicsDalhousie University Faculty of Medicine, Halifax, Nova Scotia, CanadaRichard E. BrownProfessor of PsychologyDalhousie University, Halifax, Nova Scotia, Canada in this web service Cambridge University Presswww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationUniversity Printing House, Cambridge CB2 8BS, United KingdomCambridge University Press is part of the University of Cambridge.It furthers the University’s mission by disseminating knowledge in the pursuit ofeducation, learning and research at the highest international levels of excellence.www.cambridge.orgInformation on this title: www.cambridge.org/9780521806473First edition Cambridge University Press 1994Second edition M. Wilkinson and R. E. Brown 2015This publication is in copyright. Subject to statutory exceptionand to the provisions of relevant collective licensing agreements,no reproduction of any part may take place without the writtenpermission of Cambridge University Press.First published 1994Second edition 2015Printed in the United Kingdom by Bell and Bain LtdA catalogue record for this publication is available from the British LibraryLibrary of Congress Cataloguing in Publication dataWilkinson, Michael, 1943–, author.An introduction to neuroendocrinology / Michael Wilkinson, Richard E. Brown. – Second edition.p. ; cm.Richard E. Brown’s name appears first in the previous edition.Includes bibliographical references and index.ISBN 978-0-521-80647-3 (hardback) – ISBN 978-0-521-01476-2 (paperback)I. Brown, Richard E., author. II. Title.[DNLM: 1. Neuroendocrinology – methods. 2. EndocrineGlands. 3. Neuropeptides. 4. Neurosecretory Systems. 5. Peptide Hormones.6. Receptors, Neurotransmitter. WL 105]QP356.4612.8–dc232014041712ISBN 978-0-521-80647-3 HardbackISBN 978-0-521-01476-2 PaperbackCambridge University Press has no responsibility for the persistence or accuracy ofURLs for external or third-party internet websites referred to in this publication,and does not guarantee that any content on such websites is, or will remain,accurate or appropriate. in this web service Cambridge University Presswww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationThis book is dedicated, first, to the more than 2,000 Dalhousie Universitystudents who were enrolled in the “Hormones and Behavior” undergraduatecourse and who were the original inspiration for writing the book. Many of themprovided critical comments on early drafts of the first edition.Second, one of us (M. W.) acknowledges the mentorship of the lateProfessor Kurt B. Ruf, a neuroendocrinologist and friend. in this web service Cambridge University Presswww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore information in this web service Cambridge University Presswww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationCONTENTSPreface to the second edition Acknowledgements xvList of abbreviations xvi12345xiiiClassification of chemical messengers1.11.21.31.41.51.6 1Hormones, the brain and behavior 1The body’s three communication systems 2Methods of communication between cells 5Types of chemical messenger 7Neuropeptides and neuromodulators 14Summary 16The endocrine glands and their hormones2.1 The endocrine glands 192.2 The hormones of the endocrine glands2.3 Summary 40The pituitary gland and its hormones3.13.23.33.4 45 57Functions of the hypothalamus 57Hypothalamic neurosecretory cells 58Hypothalamic hypophysiotropic hormones 62Complexities of hypothalamic-pituitary interactions Summary 73Neurotransmitters 5.15.25.35.45.519The pituitary gland 45The hormones of the pituitary gland 49Pituitary hormones in the brain 54Summary 54The hypothalamic hormones4.14.24.34.44.5 197178The neuron and the synapse 78Categories of neurotransmitters 80Neurotransmitter biosynthesis and storageRelease of neurotransmitters 92Receptors for neurotransmitters 94 in this web service Cambridge University Press 88www.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationviiiCONTENTS5.65.75.8Inactivation of neurotransmitters 97Neurotransmitter pathways 97Drugs influencing neurotransmitters, and their receptors, in thenervous system 1025.9 Can nutrients modify neurotransmitter levels and behavior? 1075.10 Divisions of the nervous system 1095.11 Summary 1146Neurotransmitter and neuropeptide control of hypothalamic,pituitary and other hormones 1206.16.26.3The cascade of chemical messengers 120Neural control of hypothalamic neurosecretory cells 120Neurotransmitter regulation of anterior pituitary hormonesecretion 1246.4 Neurotransmitter regulation of neurohypophyseal hormonesecretion 1336.5 Electrophysiology of neurosecretory cells 1356.6 Neurotransmitter regulation of other endocrine glands 1356.7 Complications in the study of neurotransmitter control ofhypothalamic hormone release 1386.8 Neuroendocrine correlates of psychiatric disorders andpsychotropic drug treatment of these disorders 1416.9 Glial cells and the regulation of hormone release 1426.10 Summary 1507Regulation of hormone synthesis, storage, release, transportand deactivation 1577.17.27.37.47.57.67.77.88The chemical structure of hormones 157Hormone synthesis 159Storage and intracellular transport of hormones 162Hormone release 163Hormone transport 164Deactivation of hormones 166Methodology for neuroendocrine research 166Summary 168Regulation of hormone levels in the bloodstream8.18.28.38.4 170Analysis of hormone levels 170Mechanisms regulating hormone levels 174Hormonal modulation of neurotransmitter release 183The cascade of chemical messengers revisited 184 in this web service Cambridge University Presswww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationCONTENTS8.58.6910When hormone regulatory mechanisms failSummary 187Steroid and thyroid hormone receptors ix1861929.19.29.3The intracellular receptor superfamily 193How are steroid hormone target cells identified? 196How are steroid hormone target cells differentiated fromnon-target cells? 1989.4 Genomic and non-genomic actions of steroid hormones 1989.5 Measurement and regulation of hormone receptor numbers 2059.6 Gonadal steroid hormone target cells in the brain 2069.7 Adrenal steroid target cells in the brain 2109.8 Steroid hormone-induced changes in neurotransmitter release 2149.9 Functions of steroid hormone modulation of nerve cells 2159.10 Thyroid hormone receptors in the brain 2239.11 Summary 225Receptors for peptide hormones, neuropeptidesand neurotransmitters 23610.110.210.310.410.510.610.7Membrane receptors 236Signal transduction by G proteins 240Second messenger systems 242Interactions in second messenger systems 248Signal amplification 249Second messengers in the brain and neuroendocrine system 249Comparison of neurotransmitter/neuropeptide and steroid hormoneactions at their target cells 25210.8 Summary 25311Neuropeptides I: classification, synthesis and co-localizationwith classical neurotransmitters 25711.1 Classification of neuropeptides 25711.2 Synthesis, storage, release and deactivation of neuropeptides 25911.3 Exploring the relationships among neuropeptides, neurotransmittersand hormones 26111.4 Coexistence (co-localization) of neurotransmittersand neuropeptides 26611.5 Localization of neuropeptide cell bodies and pathways in thebrain 26911.6 Neuropeptide receptors and second messenger systems 27211.7 Neuropeptides and the blood-brain barrier 27511.8 Summary 278 in this web service Cambridge University Presswww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationxCONTENTS1213Neuropeptides II: function286Cytokines and the interaction between the neuroendocrineand immune systems 35113.113.213.313.413.513.613.713.814 12.1 Neurotransmitter and neuromodulator actions of neuropeptides: adichotomy or a continuum? 28612.2 Neurotransmitter actions of neuropeptides 28712.3 Neuromodulator actions of neuropeptides 29312.4 Regulatory effects of neuropeptides on the neuroendocrinesystem 29912.5 Kisspeptin and GnRH as hypothalamic regulators of fertility 30012.6 Neuropeptides and the regulation of food intake and bodyweight 30212.7 Visceral, cognitive and behavioral effects of neuropeptides 31312.8 Summary 332The cells of the immune system 351The thymus gland and its hormones 354Cytokines: the messengers of the immune system 356The functions of cytokines in the immune and hematopoieticsystems 360Effects of cytokines and other immunomodulators on the brain andneuroendocrine system 364Neural and endocrine regulation of the immune system 374Hypothalamic integration of the neuroendocrine and immunesystems 384Summary 387Methods for the study of behavioral neuroendocrinology 40014.1 Behavioral bioassays 40014.2 Correlational studies of hormonal and behavioral changes 40314.3 Experimental studies I: behavioral responses to neuroendocrinemanipulation 40714.4 Experimental studies II: neuroendocrine responses toenvironmental, behavioral and cognitive stimuli 41614.5 Neural and genomic mechanisms mediating neuroendocrinebehavior interactions 42414.6 Confounding variables in behavioral neuroendocrinologyresearch 43414.7 Summary 444 in this web service Cambridge University Presswww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationCONTENTS15xiAn overview of behavioral neuroendocrinology: present, pastand future 45815.1 The aim of this book 45815.2 The history of endocrinology and behavioralneuroendocrinology 46015.3 The future of behavioral neuroendocrinologyIndex 460469 in this web service Cambridge University Presswww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore information in this web service Cambridge University Presswww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationPREFACE TO THE SECOND EDITIONIn this second edition of An Introduction to Neuroendocrinology, we have rewritten andgreatly extended the original content. The revised text includes entirely new reference listsand a complete new set of illustrations. The book reflects the many advances that haveoccurred in the study of neuroendocrinology during the past 20 years. Nevertheless, andalthough the text is based largely on modern references, our primary aim is to provide anintroductory description of mammalian neuroendocrine control systems. Several books areavailable that cover this topical and clinically relevant field, but, although valuable, thesetend to be advanced texts of the edited, multi-author type. Our book is designed to providethe basic principles necessary to understand how the brain controls, and responds to, theendocrine hormones. It will be suitable for a variety of different students and especiallythose who might not have been previously exposed to a focused course in neuroendocrinology. Thus, students in psychology, biology and science should be able to master much ofthe basic material. However, the book is also highly appropriate for honors students andfirst-year graduate students in physiology, anatomy, neuroscience and medicine. This bookis therefore designed for students in two levels of classes: introductory classes, in which allof the material will be new to the student, and more advanced classes, in which the studentswill be familiar with many of the terms and concepts through courses in biology, physiology, psychology or neuroscience, but who have not studied neuroendocrinology as anintegrated discipline.This book offers an overall outline of the neuroendocrine system and will provide thevocabulary necessary to understand the interaction between hormones and the brain. Inaddition, we provide a concise description of those topics that must underpin any attemptto learn, and to teach, neuroendocrinology. For example, there are chapters on basicneuroscience (neurotransmitters and neuropeptides), the physiology of the endocrineglands (hormones), receptors and receptor signaling mechanisms (e.g. G proteins; nuclearreceptors), hormone assay and gene expression techniques (e.g. ELISA; in situ hybridization) and a description of the immune system, with particular emphasis on the integrationof immune and neuroendocrine pathways. This basic information is also essential tounderstand the profound effects of hormones on behavior, described in Chapter 14. Oncethis material is mastered, the study of how hormones influence developmental neuralprocesses and behavior will be easier. Moreover, we have included throughout the bookreferences to the clinical relevance of many topics; for example, the influence of neuropeptides in the control of body weight and obesity. However, this book focuses primarily on theneural actions of hormones, and many of the peripheral physiological actions of hormones,such as regulation of metabolism, water balance, growth, and the regulation of calcium,sodium and potassium levels, which are the focus of traditional endocrinology texts, arereferred to only in reference to their importance in the neuroendocrine system.The introductory (second- or third-year undergraduate) student can be expected tofollow the material in this book at the level presented. To help in this, review/study in this web service Cambridge University Presswww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationxivPREFACE TO THE SECOND EDITIONquestions are given at the end of each chapter. These should be treated as practiceexamination questions and answered after each chapter is completed. For further detailedinformation on the topics covered in each chapter, all students can consult selectedreferences provided in the text. Additional references under “Further reading” are alsoincluded at the end of each chapter and these will be particularly useful to the moreadvanced student. The book will be especially relevant for more advanced (honors andgraduate) students who can use this book as an introductory account of the subject mattercovered in each chapter. These students may then take advantage of the many referencescited in each chapter to provide current and relevant information on each topic. The essayquestions at the end of each chapter also serve to provide topics for discussion, analysis anddirected research papers for the advanced student. in this web service Cambridge University Presswww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationACKNOWLEDGEMENTSThe authors are indebted to friends and colleagues who offered generous and invaluableassistance in the writing of this book. Paul Wilkinson, Ms. Alex Pincock and Ms. DianeWilkinson created several figures; Alex Pincock and Dr. Jim Pincock carefully read, andmade useful suggestions for improvement of, several early chapters. Special thanks are dueto Diane Wilkinson, who typed all the tables and assisted in compiling the extensivereference lists. The following scientists unselfishly provided illustrations from their published material: Dr. O. Almeida, Dr. A. Armario, Dr. R. Bridges, Dr. R. Goyal, Dr. L. De Groot,Dr. L. Hale, Dr. J. Herman, Drs. T. Horvath and M. Dietrich, Ms. A. Rain, Dr. T. Smith, Dr.J. Ström, Dr. J. Wakerley, Dr. A. Winokur and Dr. S. Winters. As far as we are aware, allsources of the illustrations used have been acknowledged. Permission to use previouslypublished figures was obtained either from the original authors or via RightsLink(Copyright Clearance Centre).Finally, thanks are due to Megan Waddington of Cambridge University Press for herpatience in awaiting the delivery of this manuscript. in this web service Cambridge University Presswww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore LABNPCa2 CAHcAMPthird ventricle2-arachidinoyl glycerol5-hydroxyindoleacetic acid5-hydroxytryptamine enyl cyclaseacetylcholineadrenocorticotropic hormoneantidiuretic hormone (vasopressin)attention deficit hyperactivity disorderanandamideagouti-related proteinanterior hypothalamusanterior hypothalamic areaα-amino-3-hydroxy-5-methyl-4isoxazole propionic acidamygdalaatrial natriuretic peptideautonomic nervous systemarea postremaantigen presenting cellamine precursor uptake anddecarboxylationandrogen receptorarcuate nucleusangiotensinadenosine triphosphatearginine vasopressinanteroventral periventricular nucleusβ2-adrenergic receptorβ-endorphinβ-Galactosidase immunoreactivityblood-brain barrierbrain-derived neurotrophic factorbasolateral amygdalaB-type natriuretic peptidecalcium ioncongenital adrenal hyperplasiacyclic adenosine monophosphate in this web service Cambridge University PressCARTcGMPCB1CBGCCKCCK-KOCGRPChATCLCl DHEADHTdISONDMNDMTDNAcocaine- and amphetamine-regulatedtranscriptcyclic guanosine monophosphatecannabinoid receptor 1corticosteroid binding globulin(transcortin)cholecystokininCCK knockoutscalcitonin gene related peptidecholine acetyltransferasecentrolateral thalamuschloride ioncorticotropin-like intermediate lobepeptidecentromedial thalamuscentral nervous systemcatechol o-methyl transferasecaudate/putamencAMP responsive element bindingproteincorticotropin-releasing factor (alsocalled CRH)corticotropin-releasing hormone (alsocalled CRF)cerebrospinal fluidcircumventricular organsdiestrusdopamine 2 receptordiestrus 3dopaminediacylglycerolDNA binding domaindopamine beta-hydroxylasedentate olateral supraoptic nucleusdorsomedial hypothalamic nucleusdimethyltryptaminedeoxyribonucleic acidwww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationLIST OF DPGFPGHGHRHGH-RIHGIGiGIPGLP-1GLP-2GluGM-CSFDiffuse Neuroendocrine Systemdynorphinestradiolendocrine disrupting chemicalsepidermal growth factorexternal granule cell layerejaculation latencyenzyme-linked immunosorbentassayenkephalinenteric nervous systemendogenous opioid peptideerythropoietinendoplasmic reticulumestrogen receptorestrogen response elementfibroblast growth factorfunctional magnetic ing hormonefollicle-stimulating hormonereleasing hormonefornixgranule cellsgranulocyte colony stimulatingfactorgamma-aminobutyric acidGABA transaminaseglutamic acid decarboxylaseglial-derived neurotrophic factorguanosine diphosphategreen fluorescent proteingrowth hormonegrowth hormone releasing hormonegrowth hormone release inhibitinghormone (see SOM)gastrointestinalinhibitory G proteingastrin inhibitory peptideglucagon-like peptide-1glucagon-like peptide-2glutamategranulocyte-macrophage colonystimulating factor in this web service Cambridge University LILlMANIP3iRIRS-1JAKK KPLHxviigonadotropin inhibitory hormonegonadotropin-releasing hormoneG-protein-coupled receptor 54glucocorticoid receptorglucocorticoid response elementstimulatory G proteingeneral transcription factorguanosine triphosphatehormone binding domainhuman chorionic gonadotropinhuman chorionicsomatomammotropinhistidine decarboxylasehigh fat diethepatic glucose productionhypothalamic-pituitary-adrenalhuman placental lactogenhigh performance liquidchromatographyhormone response elementhormone replacement therapyheat shock proteinhomovanillic acidnucleus intercollicularisintromission frequencyinterferon γimmunoglobulininsulin-like growth factor;somatomedininsulin-like growth factor bindingproteininternal granule cell layerinter-intromission intervalinterleukinintromission latencylateral magnocellular nucleus of theanterior nidopalliuminositol triphosphateion channelinsulin receptor substrate 1janus kinasepotassium ionkisspeptinluteinizing hormone (also lateralhypothalamus)www.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationxviiiLIST OF H-RFluteinizing hormone releasing hormonelipotropic hormone (also β-lipotropin)lysergic acid diethylamidemuscarinicmonoamine oxidasemediobasal hypothalamusmelanocortinmacrophage colony stimulating factordorsomedial thalamusmedian eminencemetestrusmossy fibersmount frequencymetabotropic glutamate receptormajor histocompatibility complex3-methoxy-4-hydroxyphenylglycolmilli international unitsmount latencymolecular layermedial geniculate bodymain olfactory epitheliummedial preoptic areametabotropic membrane receptormineralocorticoid receptormidbrain reticular formationmagnetic resonance imagingmessenger ribonucleic acidα-melanocyte-stimulating hormonemelanocyte-stimulating hormone –releasing factorMSH-RH melanocyte-stimulating hormone –releasing hormoneMSH-RIF melanocyte-stimulating hormone –release-inhibiting factorMSH-RIH melanocyte-stimulating hormone –release-inhibiting hormoneMTmelatoninMUAmultiple unit activityNAnoradrenaline (also norepinephrine, NE)Na sodium ionNEnorepinephrine (also noradrenaline,NA)NGFnerve growth factorNKnatural killer cellNKTnatural killer T cell in this web service Cambridge University rtatenitric oxidenitric oxide synthaseneurophysinneuropeptide YN-ethylmaleimide sensitive factorneurotransmitteramino terminal domainnucleus tractus solitariustracheosyringeal portion of the nucleushypoglossusolfactory bulboxytocinopioid receptor-like receptoroxytocin receptororganum vasculosum of the ne (also Purkinje cells)pituitary adenylate cyclase-activatingpolypeptideproprotein convertasephencyclidinepolymerase chain reactionphosphorylated CREBpost-ejaculatory intervalanterior periventricular nucleuspreproenkephalinpositron emission tomographyparallel fibersperifornical areaprostaglandin E2posterior hypothalamusphosphoinositide 3 kinaseprolactin releasing inhibiting factorphosphatidylinositol diphosphatepiriform cortexprotein kinase Aplacental lactogenphospholipase Cparasympathetic nervous systempreoptic areaRNA polymerasepro-opiomelanocortinwww.cambridge.org

Cambridge University Press978-0-521-80647-3 - Neuroendocrinology: Second EditionMichael Wilkinson and Richard E. BrownFrontmatterMore informationLIST OF SSTSTATTT3T4TBGTCTF5TGFαTGFβ1THTHTHCpancreatic polypeptideprogesterone receptorprolactin releasing factorprolactin-releasing hormoneprolactinproestrusprolactin-releasing peptideparathyroid hormonepost-traumatic stress disorderperiventricular nucleusparaventricular nucleuspeptide YYrobust nucleus of the arcopalliumrough endoplasmic reticulumretrosplenial cortexsubcutaneoussuprachiasmatic nucleussexually dimorphic nucleusstandard error of the meansex hormone binding globulinsoluble SNF attachment proteinsSNAP receptor proteinspinal nucleus of thebulbocavernosussympathetic nervous systemsuppressor of cytokine signalingsomatostatinsupraoptic nucleusSubstance Psomatosensory cortexsomatostatin receptorsignal transducer and activator oftranscription / signal transductionand thyroid hormone binding globulincytotoxic T cellthymosin fraction 5transforming growth factor αtransforming growth factor β1tyrosine hydroxylasehelper T celltetrahydrocannabinol in this web service Cambridge University ONVMHVMNVNOVPWATxixtuberoinfundibular DAtumor necrosis factor αthyroid hormone receptorsthyrotropin (TSH) releasing factor(also TRH)thyroid hormone releasing hormonetyrosine receptor kinasesuppressor T cellthyroid-stimulating hormoneTSH receptorthyroid-stimulating hormonereleasing hormone (TRH)vascular endothelial growth factorvasoactive intestinal polypeptideventromedial supraoptic nucleusventromedial hypothalamic nucleusventromedial nucleus ofhypothalamusvomeronasal organvasopressinwhite adipose tissue (fat)www.cambridge.org

An Introduction to Cambridge Unive rsit y Pre ss 978-0-521-80647-3 - Neuroendocrinology: Second Edition Michael Wilkinson and Richard E. Brown Frontmatter

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