Vertebrate Endocrinology, Fourth Edition

574 Pages·2006·25.63 MB·English

by  David O. Norris

Vertebrate
Endocrinology
FOURTHEDITION
This page intentionally left blank
Vertebrate
Endocrinology
F O U R T H E D I T I O N
David O. Norris
AMSTERDAM•BOSTON•HEIDELBERG•LONDON
NEWYORK•OXFORD•PARIS•SANDIEGO
SANFRANCISCO•SINGAPORE•SYDNEY•TOKYO
AcademicPressisanimprintofElsevier
ElsevierAcademicPress
30CorporateDrive,Suite400,Burlington,MA01803,USA
525BStreet,Suite1900,SanDiego,CA92101-4495,USA
84Theobald’sRoad,LondonWC1X8RR,UK
Thisbookisprintedonacid-freepaper.(cid:2)(cid:3)
Copyright©2007,ElsevierInc.Allrightsreserved.
Nopartofthispublicationmaybereproducedortransmittedinanyformorbyanymeans,
electronicormechanical,includingphotocopy,recording,oranyinformation
storageandretrievalsystem,withoutpermissioninwritingfromthepublisher.
PermissionsmaybesoughtdirectlyfromElsevier’sScience&TechnologyRights
DepartmentinOxford,UK:phone:(cid:2)+44(cid:3)1865843830,fax:(cid:2)+44(cid:3)1865853333,
E-mail:[email protected]
viatheElsevierhomepage(http://elsevier.com),byselecting“CustomerSupport”
andthen“ObtainingPermissions.”
LibraryofCongressCataloging-in-PublicationData
Norris,DavidO.
Vertebrateendocrinology/author,DavidO.Norris.—4thed.
p. cm.
Includesindex.
ISBN-13:978-0-12-088768-2(alk.paper)
ISBN-10:0-12-088768-1(alk.paper)
1. Vertebrates—Endocrinology. I. Title.
QP187.N672007
596(cid:4).0142—dc22
2006020110
BritishLibraryCataloguinginPublicationData
AcataloguerecordforthisbookisavailablefromtheBritishLibrary.
ISBN13:978-0-12-088768-2
ISBN10:0-12-088768-1
ForallinformationonallElsevierAcademicPresspublications
visitourWebsiteatwww.books.elsevier.com.
PrintedintheUnitedStatesofAmerica
07 08 09 10 9 8 7 6 5 4 3 2 1
Working together to grow
libraries in developing countries
www.elsevier.com | www.bookaid.org | www.sabre.org
This book is dedicated to my family (spouse Kay and daughters Sara Engel and Linda Quintana) who have
supported me even when I haven’t deserved it, and to the hundreds of bright, inquiring students at the
UniversityofColoradothathavemadethelast40yearsofteachingandresearchinvertebrateendocrinology
a thorough delight.
This page intentionally left blank
CONTENTS
Preface viii
1. An Overview of Chemical Bioregulation in Vertebrates 1
2. Methods to Study Bioregulation 30
3. Synthesis, Metabolism, and Actions of Bioregulators 46
4. Organization of the Mammalian Hypothalamus-Pituitary Axes 106
5. The Hypothalamus-Pituitary System in Non-Mammalian Vertebrates 168
6. The Hypothalamus-Pituitary-Thyroid (HPT) Axis of Mammals 221
7. The Hypothalamus-Pituitary-Thyroid (HPT) Axis of Non-Mammalian Vertebrates 244
8. The Mammalian Adrenal Glands: Cortical and Chromaffin Cells 272
9. Comparative Aspects of Vertebrate Adrenals 298
10. The Endocrinology of Mammalian Reproduction 322
11. Comparative Aspects of Vertebrate Reproduction 371
12. Bioregulation of Feeding, Digestion, and Metabolism 427
13. Comparative Aspects of Feeding, Digestion, and Metabolism 470
14. Bioregulation of Calcium and Phosphate Homeostasis 486
Appendix A. Abbreviations of Endocrine Terms 512
Appendix B. Vertebrate Tissue Types 519
Appendix C. Amino Acids and Their Symbols 522
Appendix D. Bioassays 523
Index 531
vii
PREFACE TO THE FOURTH EDITION
Vertebrate Endocrinology has evolved into a 4th edition that has incorporated new information and
insights gained by this author during a fourth decade of teaching and research within the field of
endocrinology. It represents a virtual rewrite and reorganization with the addition of many new
chemical bioregulators as well as new understandings of the synthesis, actions, and metabolism of
former bioregulators. Included are new insights into the evolution of these systems. Furthermore,
the old distinction of separate regulatory systems (e.g., nervous, endocrine, immune, paracrine) has
becomesoblurredduringthepastdecadethatwenowmustconsideranorganismtobeanintegrated
playingfieldforchemicalbioregulatorsproducedinavarietyofmannersbyagreatvarietyoftissues
and organs with a considerable amount of cross-talk. Perhaps, even the title of this edition should
have been altered to simply The Chemical Bioregulation of Vertebrate Physiology and Behavior to
reflect this change of emphasis from endocrinology sensu stricto.
I hope this new edition will continue the tradition of introducing undergraduate and graduate
students to the exciting, complicated, and integrative field of “endocrinology” that is more broadly
interpreted in this edition as “chemical bioregulation.” Whether interested in clinical, evolutionary,
and/or environmental aspects of endocrinology, it is my intention that students receive a broad
exposure in this book that will enable them to become self-learners by providing a background to
the primary literature.
Although tremendous gains in molecular techniques since the 3rd edition have allowed creative
scientists to greatly increase our understandings of the intricate interrelatedness among chemical
bioregulatory mechanisms and their evolution, perhaps the most important changes have been the
recognitionanddocumentationofendocrinedisruptionbychemicalsintheenvironment.Theseenvi-
ronmentally endocrine active chemicals (EACs) are capable of disrupting normal life history events
including development, sexual differentiation, and post-embryonic functions such as metabolism,
stress responses, sexual maturation, reproduction, and behavior. The possible clinical implications
of these EACs for present human populations are of immediate concern, and there is mounting
evidence of their potential impacts on future generations. Some of these EACs have appeared in
the environment simply through careless and/or thoughtless behavior related to our obsessions for
“betterlivingthroughchemistry”andfor“economicgain”regardlessofthecosttotheenvironment.
Other EACs are the consequence of the burgeoning size of the human population, its concentra-
tions into cities, and its reliance on artificially maintained agriculture and animal husbandry. EACs
include pesticides, herbicides, antibiotics, fertilizers, and industrial products (e.g., PCBs, dioxins,
and plasticizers) as well as ingredients of household products such as detergents, shampoos, and
cosmetics.Furthermore,highdensitiesofhumansanddomesticanimalshaveresultedintheaddition
of biologically relevant concentrations of reproductively active hormones and pharmaceuticals to
aquaticenvironmentsthroughwastewatereffluentsandapplicationofbiosolidstoagriculturalfields.
Of greatest long-term (evolutionary) concern perhaps may be the effects of thyroid inhibitors on
central nervous system development and the effects of estrogenic and androgenic EACs on sexual
viii
Preface ix
differentiation (sex reversal and intersex production) of embryos and immature animals as well as
the induction of estrogen-based cancers and contraception in adults and future generations. The
widespread use of pharmaceuticals, especially in the “developed” countries, is resulting in the pres-
ence of biologically relevant quantities of these drugs in water supplies and even in drinking water.
Among these drugs are estrogenic birth control ingredients, (cid:4)-blockers used for blood pressure
control or cardiac therapy, selective serotonin reuptake inhibitors (SSRIs) such as fluoxetine (i.e.,
Prozac®(cid:3), and plasma cholesterol-lowering compounds, to name a few. The widespread contami-
nation of natural environments means that it is no longer possible to study animals under natural
conditions.Hence,fieldstudiesmustrelyonreferencepopulationslackingonlythebioregulatorthey
wishtostudyratherthancontrolpopulationssinceallanimalsinnatureareexposedtosomeEACs.
And the ultimate impacts of EACs on microorganisms, plants, and invertebrate populations and
theirimplicationsfornaturalecosystemsmaybeevenmoreimportantthanthoseonvertebrates.Itis
myhopethatstudentswhounderstandtheintricaciesandinterrelatednessofchemicalbioregulatory
systems such as described in this text will be best prepared to deal with the threat of environmental
EACs in the future. If left unchecked, EACs could lead to the destruction of natural ecosystems as
we know them.
ACKNOWLEDGEMENTS
This book is in part a product of my need to teach the excitement of discovery in the biological
sciences that was instilled in me by my undergraduate advisor and botany professor at Baldwin-
Wallace College, Dr. Donald S. Dean. Were I to attempt to name all of the endocrine people who
have contributed directly or indirectly to this edition, I would undoubtedly overlook many of them.
I can acknowledge the vast majority, however, by simply referring to the many direct academic
descendents and postdoctoral students of my past mentors in vertebrate endocrinology: Aubrey
Gorbman, Howard Bern, and Donald Farner as well their many academic sons and daughters,
grandsonsandgrand-daughters,great-grandsonsandgreat-grand-daughters,etc.Thesedescendents
continue to add to our understanding of vertebrate endocrinology through teaching and research.
My own understanding of chemical bioregulation has been strongly influenced by my many years
of interactions with colleagues Dr. Richard Evan Jones and more recently Dr. Pei-San Tsai, as well
as with the many undergraduate and graduate students in vertebrate endocrinology we have trained
inourlaboratoriesattheUniversityofColoradoinBoulder.Questions,challenges,andindependent
interpretations of the literature by these students have strongly affected my thinking about chemical
bioregulation over the years to the extent that I can no longer honestly separate their ideas from
my own. Among those former graduate students are
Dr. Harriet B. Austin
Dr. James A. Carr
Dr. Laura M. Carruth
Dr. Rubai Ding
Dr. David Duvall
Dr. Kevin T. Fitzgerald
Dr. William A. Gern
Dr. John C. Gill
Dr. Louis P. Guillette, Jr.
Dr. Earl T. Larson