SECOND

REVISED

EDITION

Pathways to PREGNANCY and PARTURITION

SECOND

REVISED

EDITION

P.L. Senger, Ph. D. Emeritus Professor

Washington State University
Pullman, Washington 99164-6332 USA
Current Conceptions, lnc.

Washington State University Research & Technology Park

1610 NE East-gate 8/vd., Pullman, WA 99163-5607

Website: www.currentconceptions.com
E-mail: cci@pullman.com

Current Conceptions, lnc. 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, mechanical, recording or otherwise) without written permission
from the copyright owner.
Every effort has been made to ensure the accuracy of this work. Neither the
author nor Current Conceptions, lnc. assumes any legal responsibility or
liability for errors, omissions or method of presentation of information in
this book.

ISBN 0965764826

2'"1 Revised Edition

Phillip

L. Senger, Author

Printed in the United States of America

Communications

by: Cadmus Professional

First Edition, 1997

First Revised Edition,

1999

Second Edition, 2003

Additional copies may be ordered from:

Current Conceptions, lnc.
1610 NE East-gate Blvd. Pullman, WA 991635625
www.currentconceptions.com
Phone:
FAX:
Email:

5093345193
5093380963
cci@puUma11.con1

Cover art: Julie Steel

Cover design: Sonja Gerard Photography: Henry Moore, Jr. Prepress layout:
J. Richard Scott
A1t and Graphics: Sonja Gerard, Oei Graphics, Bellevue,
WA(soei@oeigraphics.com)
vii

Dedication

To paraphrase Hodding Carter. there are two lasting benefits we can give our
students, children and other people we associate with.

"One is roots and the other is wings.,,

This book is dedicated to Dr. R.G. (Dick) Saacke and his wife, Ann, a
couple who have been models for countless students, fellow
educators/professionals, friends and their own children in emphasizing the

importance of uncompromising commitment to high standards and values

(roots). At the same time, they have always encouraged others to fly and to
enjoy their journey (wings).

Ann Saacke and Dr. R. G. Saacke

The Author

(Phill) Senger grew up in Cary, North Carolina and received his B.S. in Zoology
from North Carolina State University. He was awarded the M.S. and Ph.D. in
reproductive physiology from the Department 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 animal physiology and reproductive physiology. He
has authored over 200 scientific, educational and popular press papers. Dr.
Senger is currently President of Current Conceptions, Inc.
Dr. Senger has 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 American Society of Animal Science
Distinguished Teaching Award in 1998 and the Marion E. Smith Faculty
Achievement Award in 2005. He has received two national awards for
research in reproductive physiology Dr. Senger has been a frequent speaker,
columnist and author about dairy reproduction to producers and veterinarians
throughout the world. He is married and has three daughters. He enjoys
racquetball, skiing, fishing, gardening and attempts golf.

Preface

The ultimate goal of pathways to Pregnancy and Parturition 2 Revised

Edition is to enable people to understand the principles of reproductive
physiology. This discipline is a visual one and requires images of anatomical
structures and physiologic processes. Good
visual imaging makes learning easier, less time consuming and the knowledge
is retained longer. Students, professors, industry professionals, veterinarians or
those simply interested in the reproductive process are very busy. They do not
have time to "confuse" themselves or others. In this context, we have tried
hard to make this book a good investment in visual understanding.
There will always be room for considerable debate concerning what is "correct"
and "incorrect" regarding many of the topics in this book. During my career as
an educator and researcher I have come to realize that,

"it is not only about making sure the information is correct ... it is a/so about
making sure people can understand it.

If they can 't understand it, it real/y doesn't matter if it is corrrect"

Dr. R.C. Mittel hammer, one of my 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 this book and we have
purposely linked them to the principles and concepts. We have not "dumped"
a plethora of facts on the reader, Instead, we have presented the concepts
(and the facts that support therm) in a logical sequence so that the reader can
easily follow the progression of key events through the reproductive process.

Inspiration

Have received numerous suggestions, criticisms and praise regarding the 2"d
Edition from university faculty, veterinarians, industry professionals and
students from all over the world. Their reactions have always been directed
toward increasing the power of this instrument to enlighten people about
reproductive physiology. Such feedback has served as a huge source of
inspiration to me and my associates during the preparation of the 2 Revised
Edition.

Blood, Sweat and Tears

Remodeling requires that a mess be made before the final product can be
visualized. Implementing the many changes, additions and modifications
has at tunes been exhausting. People with boundless energy, patience
and the relentless desire to produce quality have made huge contributions to
this edition.

Those pictured above have become known as "The Ganada/ Gossip Gong"

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 Prepress Proofing and Editing

P.L. Senger Author
Angela C. Oki Text Layout and Formatting
J. Richard Scott Photographic Presentations and Prepress Layout Chrtstina M.
Chesvick (not pictured) "Further Phenomena for Fertility" Caitlin Price (not
pictured) Index Revision
Sarah Bobbitt (not pictured) Index and Glossary Revision

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 (Emeritus, Colorado State University) L.F. ArcbbaJd (University of
Florida)
C.E. Farin (North Carolina State University) M.J. Fields (University of Florida)
D.L. Foster (University of Michigan)
fI.A. Garverick (University of Missouri) E.K. Ins keep (West Virginia University)
L. Johnson (Texas A&M University) L.S. Katz (Rutgers University)
T.L. Ott (University Idaho)
M.F. Smith (University of Missouri) M. Tatum (Texas A&M University)
A. Tibary (Washington State 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. Blair BS, Animal Sciences, Washington State University, 2002.
She assisted to the development of Figure 41 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 39 and 3- 1 O were produced as part of a Washington State University
Honors College Thesis entitled, "Intravascular Polymerization as a Method of
Observing Counter current Exchange Systems in Bovine Reproductive Tracts,"
1998. The project was sponsored by Current Conceptions, Inc., Pullman, WA.

Christina M. Davis BS, Animal Sciences, Washington State University, 2002.

Figures 154 through 158 were produced as part of a Washington State
University Honors College Thesis entitled, "A Full Color Photographic
Description of Postpartum Uterine involution in. the Dairy Cow," 2002. The
project was sponsored by Current Conceptions, ICC., Pullman, WA.
Melinda Fernyhough
BS, Animal Sciences, Washington
University, 2000. Assisted with legend layout/design.
Brian R. Voortman
University,
1998.

State

BS, Animal Sciences, Washington State

DVM, Washington State

University 2002. Assisted in the preparation of placenta! specimens and

vascular casting.
The following individuals provided valuable technical assistance.
S.R. Fenmore Radiology Technician, College of Veterinary Medicine,
Washington State University
J.M. Hobbs Clinical Lab Technician, Washington Animal Disease and
Diagnostic Laboratory
P.L. Johnson Instructional Lab Supervisor, College of Veterinary Medicine,
Washington State University
F.M. Mellieon Preliminary organization / layout
V.L. Mitzmberg Technician Supervisor, College of Veterinary Medicine,
Washington State University
T.R. Olson Scientific .Instructional Technician, College of Veterinary
Medicine, Washington State University L.M. Robinson Scientific Instructional
Technician, College of Veterinary Medicine, Washington State University B.A.
Toms Project Manager, Cadmus Professional Communications

Table of Contents

Chapter I
introduction to Reproduction
Chapter 2
The Organization and Function of the Female Reproductive Systcm
Chapter 3
The Organization and Function of the Mate Reproductive System
Chapter 4
Embryogenesis of the Pituitary Gland and the Male or Female Reproductive
System
Chapter 5
Regulation of Reproduction Nerves, Hormones and Target Tissues
Chapter 6
Puberty.
Chapter 7
Reproductive Cyclicity Terminology and Basic Concepts
Chapter 8
Reproductive Cyclicity The follicular Phase
Chapter 9
Reproductive Cyclicity The Luteal Phase
Chapter 10
Endocrinology of the Male and Spermatogenesis
Chapter 11
Reproductive

Behavior.

Chapter 12
Spermatozoa in the Female Tracer Transport, Capacitation and Fertilization
Chapter 13
Early Embryogenesis

and Maternal Recognition of Pregnancy

Chapter 14
Placentation, the Endocrinology of Gestation and Parturition
Chapter 15.

The Puerperium and Lactation

Glossary

introduction to REPRODUCTION

Reproduction is a sequence of events beginning with. development of the

reproductive system in the embryo. After the animal is born, it must grow and
achieve puberty by acquiring the ability to produce fertile gametes. This
ability must be accompanied by reproductive behavior and copulation, After
copulation, the sperm and egg meet fertilization occurs and development of
the preattachment embryo follows. The conceptus attaches to the uterus by a
specialized organ called the placenta. It allows the conceptus to grow and
develop to term. The fully developed fetus is born, and the female giving birth
to it must lactate to provide
nourishment for the neonate. During or after Lactation the dam must
reestablish cyclicity before she can become pregnant again. Knowledge and
understanding of the reproductive process will become increasingly important
as the human population continues to grow and resources become increasingly
scarce.

Welcome lo the exciting and fascinating subject of reproductive physiology,

Among the many scientific subjects in the natural sciences, knowledge about
reproductive physiology commands interest even among those who have no
scientific inclination at all. Inits broadest sense, the subject of reproductive
physiology carries with it interest, imagination, expectation, emotion and an
intrinsic desire to know more. The average person on the street could care
less about Boyle's Law, Beer's Law, the periodic table or the phylogenetic
organization of plant and animal kingdoms. But, mention copulation,
ejaculation, spermatozoa, pregnancy, the uterus, fertilization, embryo
development or any of the myriad terms associated with reproduction and
most people will be interested. Almost without exception, everyone wants to
know more about the reproductive process, whether it relates to humans, food
producing animals, their pet or just for the sake of having more knowledge.

Reproductive Physiology Consists of Several Subspecialities.

The field of reproductive physiology is a subspecialty of the physiology

discipline. In its broadest context, reproductive physiology can be defined as
the study of reproduction in animals regardless of species. In the field of

Animal Sciences, reproductive physiology is a general term used to describe

a field of study that deals primarily with reproduction in food producing
animals. The terms andrology, gynecology, theriogenology and obstetrics a II
imply a clinical application associated with reproductive function in humans
and animals. Andrology is a branch of reproductive physiology that deals
specifically with the study and treatment of male animals including humans.
Gynecology is a branch of reproductive physiology and medicine that deals
specifically with reproductive issues in women. Theriogenology is a branch of
veterinary medicine that focuses on the reproductive system in animals.
Obstetrics is a branch of reproductive physiology, veterinary medicine and/or
human medicine that specializes in the female before, during and after
parturition.

There is a wide breadth of opportunities in the field of reproductive physiology

that range .from animal production, clinical, educational and research
applications. Also, pharmaceutical companies manufacture and market
hundreds of hormones and drugs that manipulate reproductive functions in
humans and animals. These corporations provide significant research,
technology transfer and marketing opportunities for those with training in
reproductive physiology.

Introduction to Reproduction

Figure 1-1. Lifetime Sequence of Reproductive Events

Puberty

After puberty, the female enters a period of cyclicity (C) in which repeated
estrous or menstrual cycles occur. During the cycle, copulation between
the male and female takes place.
Copulation causes pregnancy (preg). At the end of gestation the female
gives birth and enters the puerperium (P), a period of uterine and ovarian
"recovery" and begins to lactate (L). These events are repeated throughout
the female's lifetime until she enters reproductive senescence.
Successful reproduction is an orderly sequence of events. The major events
of the reproductive process are puberty, cyclicity, copulation, pregnancy and
postpartum recovery/lactation.

Figure 11 illustrates the overall lifetime sequence of these reproductive

events. As you can see from the figure, by far the majority of the
"reproductive budget'' is associated with the female.

Pathways to Pregnancy_ and Parturition is intended to help you develop a solid

scientific understanding of the principles of reproduction in domestic animals.
Further, it is intended to help you be come fluent in the language of the
subject matter. If you develop this fluency, you will enjoy a lifetime of
understanding that will enable you to adapt successfully to new knowledge
and technology that will affect reproduction in animals as well as humans.
As you use Pathways to Pregnancy and Parturition. you will encounter a
"Sequence Map" at the beginning of each chapter (See Figure 12).
In the
"sequence map," each major event is represented by a sphere positioned along
the pathway. A sign, reading "You Are Here" lets you know exactly where the
chapter you are about to read fits in the overall sequence of reproductive
events. Each event in the "sequence map" has one or more chapters
dedicated to it, These events are described briefly in the following pages.
As you read the chapters in this book you will encounter several features that
are intended to make learning and understanding easy. The text of each
chapter begins with a "Take Home Message". This feature provides you with
the main points of the chapter before you engage the details. The "Take Home
Message" should establish some questions in your mind that will then be
answered later in the chapter, The "Take Home Message" is also intended as a
study guide, highlighting the main points of each chapter.
Fact Boxes are included throughout each chapter to give you a "quick read,"
to highlight important points, terms and/or sequences and to allow you to
regroup your thoughts as you read the text.

Many words and terms in this textbook are in bold print. They are important
key words. You should understand them, know how to pronounce them, know
how to spell them and be able to use them correctly in a discussion or in
writing. In addition to the explanations appearing in the text, these terms are
also defined in the glossary at the end of the book.
At the end of each chapter is a short section called "Key References".
Important sources containing additional, in depth information about the
subject of the chapter are provided. In general, these are scientific review
papers that will provide detail beyond what is presented in the chapter.
There are some remarkable reproductive phenomena throughout the animal
kingdom. The section entitled "Further Phenomena for Fertility"(see Key
References) is intended to present some of the interesting facts, observations

and even myths relating to the topic of each chapter. This section will give
species other than domestic animals a place to shine.

Figure 1-2. Sequence Map of Reproductive Events

The male and female have a common sequence of developmental events until
after copulation. After copulation the female bears the responsibility for
gestation, parturition, lactation and post- partum uterine repair. The arrow on
the male pathway indicates his departure from the sequence after copulation.
The sign "You are here" indicates where the chapter you are about to read fits
in the sequence of reproductive events.

The Puerperium and Lactation

Parturition
Fetal Attachment and Gestation
Early Embryogenesis and Maternal Recognition of Pregnancy
Ovulation and Fertilization
Copulation
Cyclicity
Regulation of Reproduction
Spermatogenesis
Regulation of Reproduction
Tract Function
Puberty
Prenatal Development

Introduction to Reproduction
"Prenatal Development" (Chapter 4)