Table Of ContentSECOND REVISED EDITION
•
Pathways t o
P R E G N A N C Y
and
P A R T U R I T I O N
SE C OND REV I S E D EDITION
P.L. Senger, Ph. D.
Emeritus Professor
Washington State University
Pullman, Washington 99164-6332 USA
Current
Coneeptions, lne.
Washington State University Research & Technology Park
1610 NE Eastgate Blvd., Pullman, WA 99163-5607
Website: www.currentconceptions.com
E-mail: [email protected]
V
© Current Conceptions, Inc. 2005, 2003, 1999, 1997.
All rights reserved. This book is protected by copyright. No part of this book may be reproduced in any form or
by any means, including photocopying, stored in a retrieval system, transmitted in any form (electronic, mechani
cal, recording or otherwise) without written permission from the copyright owner.
Every effort has been made to ensure the accuracy of thjs work. Neither the author nor Current Conceptions, Inc.
assumes any legal responsibility or liability for errors, om.issions or method of presentation of infonnation in th.is
book.
ISBN 0-9657648-2-6
2'"1 Revised Edition
Phillip L. Senger, Author
Printed in the United States of America by: Cadmus Professional Con1munications
First Edition, 1997
First Revised Edition, 1999
Second Edition, 2003
Additional copies may be ordered from:
Current Conceptions, Inc.
1610 NE Eastgate Blvd. Phone: 509-334-5193
Pullman, WA 99163-5625 FAX: 509-338-0963
www.currentconceptions.com Email: [email protected]
Cover art: Julie Steel
Cover design: Sonja Gerard
Photograp11y: Henry ~Moore, Jr.
Pre-press layout: J. Richard Scott
A1t and Graphics: Sonja Gerard, Oei Graphics, Bellevue, WA([email protected])
vii
Dedication
-- : I • • 1 • • • ,- • ~ • • - • : • • , • • ., • ,- , ·••,# ~ , - • - • • " - _ • •
To paraphrase Hodding Carter, there are two lasting benefits
can give our students, children and other people associate with.
we we
-
"One is roots and the other is wings.''
his book is dedicated to Dr. R.G. (Djck) Saacke and his wife, Ann, a couple who
have been 1nodels for countless students, feUow educators/professionals, friends and
their own children in emphasizing the i1nportance of uncomprmnising c01n1nit111ent to
high standards and values (roots). At the same tune, they have always encouraged
others to fly and to enjoy their journey (wings).
Ann Saacke and Dr. R. G. Saacke
viii
The Author
--
- ...:~ - .
j
(Phjl) Senger grew up in Cary, North Carolina and received his B.S. in Zoology from
North Carolina State University. He was awarded tbe M.S. and Ph.D. in reproductive pl1ysiology
fro1n the Departtuent of Dairy Science at Virginia Polytechnic Institute and State University.
He has been Professor of Animal Sciences at Washington State University where his primary
teaching responsibilities include anunal physiology and reproductive physiology. He has authored
over 200 scientific, educational and popular press papers. Dr. Senger is cun-ently President of
CtuTent Conceptions, Inc.
Dr. Senger bas over 30 years of experience conducting research and teaching students, and clientele
about reproductive physiology: He is the recipient of six teaching awards, including the A,nerican
Society of Ani,nal Science Distinguished Teaching Award in 1998 and tbe Ma1ion E. Sn1ith
Faculty Ach:ieve111ent Award in 2005. He bas received two national awards for research in
reproductive physiology. Dr. Senger has been a frequent speaker, colun,nist and author about dairy
reproduction to producers and vete1inarians tlu·oughout the world. He is married and has three
daughters. He enjoys racquetball, skiing, fishing, gardening and atte1npts golf.
IX
Preface
he ultimate goal of £.athways to Pregnaocy_a,nd Parturition - 2.nd Revised Edition is
to enable people to understand tbe principles of reproductive physiology. This discipline is a
visual one and requires images of anaton1ical structures and physiologic processes. Good
visual i111aging makes learning easier, less time consun1ing and the knowledge is retained longer.
Students, professors, industry professionals, veteriJ1arians or those sitnply interested in the
reproductive process are veiy busy. They do not bave time to "deconfuse" themselves or others.
ln this context, we have tried hard to nJake this book a good investment in visual understaoding.
There will always be room for considerable debate concerning what is "correct" and "incorrect"
regarding n1any of the topics in this book. During 1ny career as au educator and researcher I have
co1ne to realize that,
"ft is not only about making sure the information is correct ...
it is also about making sure people can understand it.
If they can't understand it, it really doesn't matter if it is correct."
Dr. R.C. Mittell1a1ntner, Olle of rny colleagues at Washington State University and a renowned
educator in the field of econometrics once commented "classes, textbooks and lectures should
be much more than information dumps." Many details and facts are presented in tbis book and
we l1ave purposely linked them to the principles and concepts. We have not "dutnped" a plethora
of facts on the reader. Instead, v.1e have presented the concepts (and the facts that support tbe111) in
a logical sequence so that the reader can easily follow the progression of key events through the
reproductive process.
X
Inspiration
have received nun1erous suggestions, criticis1us and praise regarding the 2nd Edition fron1
university faculty, veterinarians, industiy professionals and students :fro1n all over the world. Their
reactions have ahvays been directed toward increasing the power of this instnunent to enlighten
people about reproductive physiology. Such feedback has served as a huge sotu-ce of inspiration
to rne and my associates dtu·ing the preparation of the 2nd Reyjsed Edition.
xi
Blood, Sweat and Tears
roduction of the 2nd Edition and the 2nd Revised Edition of Path,vay_s_to Pregnancy and
farturition has been more challenging than the 1st Edition. Remodeling requires that a mess be
n1ade before the final product can be visualized. Imple1nenting the 1nany changes, additions and
n1odifications has at ti.tnes been exhausting. People with boundless energy, patience and the
relentless desire to produce quality have n1ade huge conttibutions to tbjs edition.
Those pictured above have become known as "The Gonadal Gossip Gang"
Standing (left to right):
Henry Moore, Jr. - Photography
Dessa M. Dal Porto - Glossary
Sonja Gerard -Art, Graphics and Product Design
.Kneeling (left to right):
Renee C. Anderson - Pre-press Proofing and Editing
P.L. Senger - Author
Angela C. Oki -Tex't Layout and Formatting
J. Richard Scott - Photographic Presentations and Pre-press Layout
Christina M. Chesvick (not pictured) - "Fmther Phenomena for Fertility"
Caitlin Price (not pictured) - Index Revision
Sarah Bobbitt (not pictured) - Index and Glossary Revision
xii
Knowledge and Know-How
The following scientists made valuable contributions to the scientific content
of one or more chapters in this book.
R.P. Amann (Emelitus, Colorado State University) L. Johnson (Texas A&M University)
L.F. Archbald (University of Florida) L.S. Katz (Rutgers University)
C.E. Farin (North C,u·olina State University) T.L. Ott (University of Idaho)
M.J. Fields (University of Florida) M.F. Sn1ith (University ofMissoLtri)
D.L. Foster (University of Michigan) M . Tatum (Texas A&M University)
fl.A. Garverick (University of Missouri) A. Tibary (Washington State University)
E.K. Inskeep (West Virginia University)
During the development of 2nd Edition, a number of Baccalaureate and Veterinary students
contributed significantly. These individuals and their contributions are presented below.
Cerissa K. Blah- - BS, Animal Sciences, Washington State University, 2002.
She assisted in the development of Figure 4-1 l (inguinal hernia) in cooperation with the Washington State University Student
Swine Cooperative.
Rebecca L. Cody - BS, Animal Sciences, Washington State University, 1998, DVM, Washington State
University 2002.
Figures 3-9 and 3-10 were produced as parL of a Washington State University Honors ColJege Thesis entitled, "Intrav ascular
Poly1nerization as a Method of Observing Countercurrent Exchange Systems in Bovine Reproductive Tracts," 1998. The
project was sponsored by Current Conceptions, Inc., Pullman, WA.
Christi.ua M. Davis - BS, Anin1al Sciences, Washington State University, 2002.
Figures 15-4 through 15-8 were produced as part of a Washington State University Honors College Thesis entitled, "A Full
Color Photographic Description of Postparhun Uterine Involution in. the Dairy Cow,'' 2002. The project was sponsored by
Current Conceptions, I11c., Pullman, WA.
Melinda Fernyhough - BS, Animal Sciences, Washington State University, 2000. Assisted witb legend
layout/design.
Brian R. Voortman - BS, Animal Sciences, Washington State University, 1998. DVM, Washington State
University 2002. Assisted in the preparation of placental specimens and vascular casting.
The following individuals provided valuable technical assistance.
S.R. Feuimore - Radiology Technician, College of Veterinary Medicine, Washington State University
J.M. Hobbs - Clinical Lab Technician, Washington Animal Disease and Diagnostic Laboratory
P.L. Johuson - Instructional Lab Supervisor, College of Veterinary Medicine, Washington State University
F.M. MelUeon - Preli n1inary organization/ layout
V.L. Mitzimberg -Technician Supervisor, College ofVeterirnu·y Medicine, Washington State University
T.R. Olson - Scientific Instructional Tech11ician, College of Veterinary Med.icine, Washington State University
L.M. Robinson -Scientific Instructional Technician, College of Vete1inary Medicine, Washington State University
B.A. Toms -Project Manager, Cadmus Professional Communications
Table of Contents
Chapter I
Introduction to Reproduction . . . . .... . . .. . . . . ...... . ... . .. • ............... P.
Chapter 2
The Organization and F1111ction of the Fen1ale Reproductive Syste1n .... . .......... P. I 0
Chapter 3
The Organization and function of the Male Reproductive Syste1n . .. ........ . ..... P. 44
Chapter 4
Embryogencsis of the Pituitary Gland and the Male or F'emale Reproductive Systen1 . .. P. 80
Chapter S
Regulation of Reproduction - Nerves, J{ormones and Target Tissues . . . ...... • .... . P. I 02
Chapter 6
Pttberty .. ...... . . . ... . .. . . . .. . . . . . . . . . . . . . . . . • . . . . . . . . .. . . . . • . . . . . . . . P. I 28
~
Chapter 7
Reproductive Cyclicity - Tenuinology and Basic Concepts . . . . . . . . . . . . . . . • . . . . . . P. 144
Chapter 8
Reproductive Cyclicit.y - The follicular Phase . ..... . • ...... • ..... . ........... P. 164
_ Chapter 9
Reproductive Cyclicity - The Luteal Phase .. . . . . • .. , ..... . ........... . ...... . P. 188
Chapter 10
Endocrinology of the Male ru1d Spennatogenesis . . . . .... , ... . .. • ... . .... . ... . . P. 214
Chapter 11
Reproductive Behavior . . .. . . . . ........ • .. • .... . . . .... . . • ... • .... , .•..... P. 240
Chapter 12
Spermatozoa in the Female Tract - Transport, Capacitation and Fertilization ......... P. 266
Chapter 13
Earl)7 En1bryogcncsis and Maternal Recognition of Pregnancy .. ... . . . ...... . ... .. P. 284
Chapter 14
Placenta1ion, the Endocrinology of Gestation and Part11rition .. ... • :-......... • .... P. 304
_ Chapter IS
The Puerperiun1 aad Lactation ....... . • ... . . . . .... . . . . . • ...... •• . .• ....... P. 326
Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. 347
Index . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. 367
'