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Commissioning Editor: Robert Edwards
Development Editor: Louisa Welch
Project Manager: Jane Dingwall
Designer: Clwrles Gray
Illustrator: Oxford Illustrators
Ill ustration Managers: Kirsteen Wright and Gillian Richards
Veterinar

NINTH EDITION !

Edited by

David E. Noakes BVetMed, PhD, DSc, DVRep, DipECAR, FRCVS

70, Whitney Drive, Stevenage, Herts., UK

Timothy J. Parkinson BVSc, DBR, Dip ECAR, MEd, PhD, FRCVS

Institute of Veterinary, Animal & Biomedical Science, Massey University, Palmerston North, New Zealand

and

Gary C.W. England BVetMed, PhD, DVetMed, DVR, DVRep, DipECAR, DipACT, FRCVS
School of Veterinary Medicine & Science, University of Nottingham, Sutton Bonington Campus,
Loughboroug h, UK

SAUNDERS

ELSEVIER
Edinbur.9h London New York Oxford Philadelphia St Louis Sydney Toronto 2009
SAUNDERS
ELSEVIER
An Imprint of Elsevier Ltd.
Elsevier Limited 2009
<D

The rigllls of David Noakes, Timothy Parkinson and Cary England to be identiYied as authors of this work have
been asserted by them in accordance with the Copyright, Designs and Pate nlS Act 1988.
No pan of this publication may be reproduced or transmiued in any form or by any means, electronic or medtanical,
including photocopying, recording, o r any information storage and retrieval system, without permission in writing
from the publisher. Permissions may be sought d irectly from ElsL-vier's RigltlS Department: phone: (+I) 215 239 3804
(US) or (+44) 1865 843830 (UK); f.1X: ( +44) 1865 853333; e-mail: hea [email protected]. You may also
complete your request on-line via the Elsevier website at http:ffwww.elsevier.comfpermission.~.
First published 1938 as VctcriiiiiT}' Obstetrics byE Benesch
Second edition 1951 as VercrinaT}' Obstetrics byE Benesch and J.C. Wright
Third edition 1964 as \"/right's \TeteriiiiiiJ' Obstetrics by C.l-1. Arthur
Fourth edition 1975 as Veterinary llcfJroduction and Obstetrics by C. H. Arthur
Fifth edition 1982 as VeterinaiJ' Reproduction ami Obstetrics by C. H. Arthur, D. E. Noakes and H . Pearson
Sixth edition 1989 as Vercrinmy Reprocluctiou ami Obstetrics by C.H. Arthur, D.E. Noakes and H . Pearson
Seventh edition 1996 as Veterinary Reproduction ami OIJSierrics by C.t-1. Arthur, D.E. Noal\es, t-1. Pearson a nd
T.J. Parkinson
Eighth edition 2001 as Arthur's VeteriiUIIJ' Reproduction lllld Obstetrics by D. E. Noakes, '1:1. Parkinson and C.C.W.
England
Ninth edition 2009 as VeterillliiJ' RcprocluCiion arul Obstetrics by D.E. Noakes, T.). Parkinson and C .C.W.
England

ISBN: 978-0-7020-2887- 8

British Library Cataloguing in Publication Data

A catalogue record for this book is available from the British Library

Library of Congress Cataloging in Publication Data

t\ catalog record for this book is available from the Library of Congress

Notice
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Con tributors ..... . ...... .. .. ......... .. vii
Part Three: Dystocia and other
Preface ................................. ix disorders associated with parturition

8. General considerations .... . ....... .. 209

David Noalles
Part One: Normal cyclical ovarian
9. The approach to an obstetric case ..... 223
activity and its control
David Notziles
10. Maternal dystocia: causes and
1. Endogenous and exogenous control
treatment ...................... 232
of ovarian cyclicity . .................. 3 David Noalles
David Noalzes 11. Fetal dystocia: aetiology, incidence
and prevention .................... 247
Part Two: Pregnancy and Da11id Notzlles
parturition 12. Manipulative delivery per vaginam
in farm animals and horses .......... 266
2. Development of the conceptus . . . . . . . . 61 Dazlid Noalles
David Noalzes 13. Vaginal manipulations and delivery
3. Pregnancy and its diagnosis ...... . .... 76 in the bitch and queen cat . .......... 275
Marcel Taveme aud David Noalles David Noalles
4. Abnormal development of the 14. Dystocia due to fetomaternal
conceptus and its consequences .... . . 123 disproportion: treatment ............ 280
Susmz Long David Noahes
5. Prolapse of the cervix and vagina ..... 146 15. Dystocia due to postural defects:
David Noalles treatment. ........................ 286
6. Parturition and the care of parturient David Noalles
animals, including the newborn ...... 154 16. Dystocia due to faulty position and
Marcel Taverue and David Noalws presentation, twins and fetal
7. The puerperium ............ . ...... 194 monsters .............. . ........ . . 297
David Noalzes Dtwid Noalles

v
 . ' ..... __.,.~~-:.~~,·
~ . . .·~ : Contents
...; : .... •• l ~ t..tl... ....
..

17. Injuries and diseases incidental to Part Six: The male animal
parturition ..... ........ ........... 306 · _..#'
David Noalws 29. Normal reproduction in male
18. Postparturient prolapse of the uterus .. 319 animals .... ................ ...... 681
David Noahes T,im Parldnsotz
19. Fetotomy ......................... 326 30. Fertility, subfertility and infertility
]os Venmmt in male animals ................... 705
Tim Parhinson
Part Four: Operative interventions 31. Artificial insemination ..... .... . .... 765
Tim Parhinson
20. The caesarean operation and the
surgical preparation of teaser males ... 34 7 Part Seven: Exotic species
]os Venmmt and David Noalws
21. Genital surgery in the bitch 32. Reproduction in Camelidae .......... 809
and queen ........................ 3 7 6 Marzooh Al-Eimah
Gm1' England 33. Reproduction in the buffalo ..... .... 824
Nazir Alzmad aud
Part Five: Subfertility and infertility David Noalzes
34. Normal reproduction, reproductive
22. Infertility and subfertility in the cow: disease and infertility in pet small
structural and functional abnormalities, mammals .......... . ............. 83 6
management deficiencies and Slzaron Redrobe
non-specific infections .............. 393
Tim Parllitzson Part Eight: Assisted reproduction
23. Specific infectious diseases causing
infertility and sub fertility in cattle ..... 4 76 35. Assisted reproductive technologies .... 855
Tim ParhiliSOil Ingrid Briiclz Boglz and Torben Greve
24. Veterina1y control of herd fertility ..... 517
Tim Parlziuso11 and David Barrett
Appendix
25. Infertility and subfertility in the ewe
Hormones, related substances and vaccines
and doe (female goat) .............. 559
used in reproduction .... . ............... 895
Keitlz Smitlz
David Noalzes
26. Infenility and subfertility in the mare .. 582
Dale Paccamonti and
Jonatlum Pycoclz Index .......... . . . ....... . ........ . .. 905
27. Infertility and subfertility in the gilt
and sow .......................... 632
Olli Pelto11iemi ami Bas Kemp
28. Infertility and subfertility in the bitch
and queen .... .... .. . ............. 646
Gm1' England

VI
Professor Nazir Ahmad Professor Bas Kemp
Professor of Animal Reproduction, Professor of Adaptatio n Physiology,
DepartmeiH of Animal Reproduction, Department of Animal Sciences,
University of Agriculture, Wageningen University,
Faisalabad, Pakistan Wageningen, Netherlands

Professor Marzook AI-Eknah Dr Susan E. Long

Pro fessor ofTheriogenology, Honorary Senior Lecturer,
College ofVeterinary Medicine and Animal nesources, Department of Veterinary Clinic<JI Sciences,
King Faisal University, Al-Ahsa, · University of Bristo l,
Kingdom of Saudi Arabia Clarendon Veterinary Centre,
Weston-super-Mnre, UK
Mr David C. Barrett
Senior Lecturer in Farm Animal Health, Professor David E. Noakes
Division of Veterinary Animal Production and Public Health, Professor Emeritus of Veterinary Obstetrics and
University o f Glasgow Veterinary School. Diseases of neproduction,
Glasgow, UK Royal Veterinary College,
University of Londo n, Lo ndon, UK;
Professor Ingrid Bruck Begh Special Professor o f Veterinary neproduction,
Professor ofVeterinmy Reproduction and Obstetrics, University of Nottingham,
Department of Large Animal Sciences, Loughborough, UK
Facully of Life Sciences,
University of Copenhagen, Professor Dale L. Paccamonti
Fredericksberg, Denmark Professor and Head,
Department of Veterinary Clinical Sciences,
Professor Gary C.W. England Schoo l of Veterinary lvledicine,
Founding Dean and Professor of Veterinary neproduct.ion, Louisiana State University, Baton Rouge, l.A USA
School of Veterinary Medicine and Science,
University of Nottingham, Professor Timothy J. Parkinson
Loughborough, UK Professor of r=arm Animal Reproduction and Health,
Institute ofVeterinary, Animal and Biomedical Science,
Professor Torben Greve Massey University, Palmersto n North, New Zealand
Professor of Domestic Animal Reproductio n,
Department of Large Anim<Jl Sciences,
Faculty of Life Sciences,
University of Copenh<~gen,
FredericJ,sberg, Denmark

vii
 Contributors

Professor Olli Peltoniemi Dr Keith C. Smith

Adjunct Professor of Domestic Animal Reproduction, Tiverton, Dev~r;j)K
Departm ent of Production Animal Medicine,
University of Helsinki, Professor Marcel A.M. laverne
Saarentaus, Fin land Distinguished Pro fessor of Fetal and Perin atal Biology,
Deparupent of Farm Animal Health,
Dr Jonathan F. Pycock Faculty of Veterinary Medicine,
Director, Utrecht University,
Equine Reproductive Services, Malton, Utrecht, The Netherlands
North Yorl<shire, UK
Dr Jos J. Vermunt
Ms Sharon P. Redrobe Registered Veterinary Specialist in Cattle Med icine,
Honorary Senior Lecturer, Wellingto n, New Zea land
Department of Veterinary Clinical Sciences,
University of Bristol;
Head o f Veterinary Services,
flristol Zoo Gardens,
Bristol, UK

viii
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 It is with a great sense of pleasure, and relief, that the ninth edition of Veterinary Reproductio11 and Obstetrics
has been completed. It is sad that the late Professor Geoffrey Arthur did not live to see its completion; he
died in March 2007 having just celebrated his 9lst birthday. His contributions to the subject during his
nearly 70 years as a veterinarian, both as a stimulating teacher and mentor and as a pioneer in veterina1y
clinical research, were immense. We dedicate this book to his memory.
It is perhaps fitting that one of the features of this ninth edition is that more of its contributors are from
outside the UK than in previous editions; we have tried to 'internationalize' the contributors, who are from
eight different countries. The conception of this textbook was based on the translation into English, in 1938
under the watchful eye of the late Professor John George Wright, of what was considered at the time to be the
definitive text in the subject: Geburtsliilfe bei Rind wui Pferd by Professor Franz Benesch, who was Professor of
Obstetrics and Director of the Obstetrical Clinic in the Veterinary High School in Vienna. Subsequent!)', ).G.
Wright became co-author of the book entitled Veterinary 0/Jsletrics, which was very much a true description
of the book, since, in the first edition published in 1951 under the authorship of Benesch and Wright, 55%
of the 455 pages was devoted to classical veterinaty obstetrics. Despite many changes, we have still retained
some of the original diagrams. In the first edition there were some excellent drawings of various fetal dispo-
sitions by the late Mr A.C. Shuttleworth, who was Senior Lecturer in Veterinary Anatomy at the University
of Liverpool under Professor J.G. Wright. Initially in half-tone and meticulously drawn from postmortem
specimens, they have been copied and modified in many other textbooks. A.C. Shuttleworth was a very fine
artist, and it is fitting to recognize his legacy as an illustrator. In addition, we have also retained many of the
photographs used by the late Professors Geoffrey Arthur and Harold Pearson in previous edi tions.
It is our intention that this new edition should primarily meet the needs of the veterinary undergraduate,
although no doubt it will be useful for others. All the chapters have been updated and in some cases com-
pletely revised and rewritten by ne\v authors. Chapters 3 and 6, on pregnancy and parturition, respectively,
have had major input from Marcel Taverne, Chapter 20 on the caesarean operation in farm animals and horses
has had a major revision with substantial input from Jos Vennunt, who has also written a separate chapter
on the rapidly disappearing, but very useful, art of fetotomy. In the first editio n of the book, published in
1951, 56 out of a total of 455 pages were devoted to this topic. New illustrations for these chapters, drawn by
Peter Parkinson, have completed the renewal of this section. The chapter on porcine infertility and subfer-
tility has been completely rev.rritten by O lli Peltoniemi a nd Bas Kemp, as has Chapter 34, which deals with
normal reproduction and reproductive disease in small pet mamm als, by Sharon Redrobe. Dale Paccamonti
has collaborated with Jonathan Pycock on infertility and subfertility in the mare, as has David Barrett with
Tim Parkinson on the chapter on veterinary control of herd fertility. Finally, the list of new authors includes
Ingrid Bruck B0gh and Torben Greve, who have written a new chapter on assisted reproductive technologies,
which has greatly expanded the original chapter in the eighth edi tion on embryo transfer.

ix
.,. :' - . '!
~·· _{C-
Preface

Finally we must thank the production team at Elsevier for all their help during the preparation of this
new edition. Changes in technology have enabled. most of the illustrations to be reproduced in colour,
which has required substan tial 'shepherding' of the authors; we are.pa/acularly grateful to Louisa Welch, the
Develo pment Editor, Jane Dingwall, Project Ma nager, Sulde Hunter, Copy Editor and many o thers for their
patience and courtesy in this regard. ·

April 2009 David E. Noakes

Timothy J. Parkinson
Gary C.W. England

X
 I1 I
Normal cyclical ovarian
activity and its control

1 Endogenous and exogenous control of

ovarian cyclicity ............ ............................. 3
David Noakes
 .c·!
..Mr'
 I1 I David Noakes

Endogenous and exogenous control of

ovarian cyclicity
In nature, it is th e general rule that animals breed sexually mature and able to reproduce is referred to
once annually and pa nurition occurs in the spring, as puberty. Among females of the domestic species,
the time most favourable to the progeny, since puberty precedes the development of physical matu-
the early neonatal period of their life will be dur- rity a nd, although they become capable of repro-
ing the period of increasing light and warmth, and ducing, their efficiency, panicularly with respect to
also at the tim e when food fo r the mother is most their fecundity, has not reached its maximum.
abundant to ensure adeq uate lactation. Under The initiation of puberty is largely a function of
the conditions of feeding a nd h ousing provided the animal's age and maturity since the female is
by domestication the breeding season tends to be born with a genetic potential for cyclic reproduc-
lengthened, and some of our species, panicularly tive activity. Provided the environmental influences
cattle, may breed at any time during th e year; all are favourable at this time, then once the 'biologi-
do mesticated animals, however, show a co nstant cal clock' is started it will continue for as long as
tendency to revert to the natural breeding season, the environ ment remains favo urable. In none of
as evidenced by reduced fertility during summer our domesti c species is there a physiological change
and early autumn in sows. comparable with the m enopause of women.
For an ani mal to breed, it must be mated and hence Among non-seasonal polycyclic animals, such
must attract th e male and be sexually receptive (in as the cow and sow, the recurring cyclic activity is
heat or in oestrus). All domestic species show recur- intermpted by pregnancy, lactation and pathologi-
ring periods of sexual receptivity, or oestrous cycles, cal conditio ns. In those species that are seasonally
which are associated with the ripening in the ovaries polycycl ic, the mare, ewe, doe (or nanny) goat and
of one or more graafian follicles (Fig. 1.1) and cul- cat, or monocyclic, such as the bitch, there are peri-
minate in the sheddi ng of one or more ova. If a fer- ods of sexual quiescence or a noestms.
tile mating occurs then pregnancy may ensue. When the fe male reaches puberty the geni-
tal organs increase in size. During the prepubertal
period the growth of the genital organs is very sim-
Puberty and the onset of cyclic
ilar to that o f o ther organ systems, but at puberty
activity their growth rate is accelerated, a point well illus-
,• trated in the gilt, where the mean length of the uter-
The young fe m ale animal shows no evidence of ine horns is increased by 58%, the m ean weight
recurring or cyclic periods of sexual receptivi ty. The of the utems by 72% and the m ean weight of the
onset of such changes when the female becomes ovaries by 32% between 169 and 186 days of age

3
 I 1 I Normal cyclical ovarian acti:vity and its control

there were follicular waves in response to follicle-

stimulating hormone (FSI-1) secretion that were simi-
lar to those of--tft'e adult, and that individual follicular
development was characterized by growing, static and
regressing phases (Adams 1994).
The sheep has been used extensively for studying
many of the mechanisms involved in the initiation
of puberty; however, it must be stressed that season-
ality will exert an overriding influence in this spe-
cies (see below). The onset of puberty is signalled
by either the occurrence of the first oestrus or the
first ovulation; in the ewe lamb these do not occur
simultaneously because the first ovulation is not
preceded by behavioural oestrus. A similar response
Fig. 1.1 Cross-section of a graafian follicle.
is seen in sexually mature ewes at the onset of the
normal breeding season.
(Lasley 1968). Females of domestic species reach the The hormone that is primarily responsible for
age of puberty at the following times: the onset of ovarian activity, and hence puberty, is
• mare: 1-2 years luteinizing hormone (LI-1). In adult ewes during the
normal breeding season, basal Ll-1 concentrations
• cow: 7-18 months
increase together with the LH pulse frequency to one
• ewe: 6-15 months per hour during the period of maximum follicular
• doe or nanny goat: 4-8 months growth. This results in the development of follicles to
• sow: 6-8 months the preovulatory stage, and their secretion of oestra-
diol, which activates the LI-1 surge causing ovulation
• bitch: 6-20 months
and corpus luteum formation. In the prepubertal
• queen cat: 7-12 months. ewe Iamb, LI-1 pulses occur at similar amplitudes but
The changes that occur at puberty depend directly much lower frequencies (one every 2-3 hours). As a
upon the activity of the ovaries, which have two consequence, follicular growth is insufficient to acti-
functions: the production of the female gametes vate the LI-I surge necessary for final follicular matu-
and the synthesis of hormones. Let us consider the ration and ovulation.
changes that occur in the ovary of the young heifer Experimental evidence in prepubertal ewe lambs
calf. At birth, each ovary may contain up to 150 000 has shown that ovarian follicles are capable of
primary or primordial follicles; each consists of an responding to exogenous gonadotrophin stimula-
oocyte surrounded by a single layer of epithelial tion and the pituitary is capable of secreting Ll-1 at
cells but with no thecal cells. Soon after birth, the a frequency to stimulate ovulation. The failure of
ovaries start to develop and produce growing folli- the prepubertal ewe lamb to undergo ovulation and
cles, which consist of an oocyte with two or more exhibit oestrus is due to the high threshold for the
layers of granulosa cells and a basement membrane. positive-feedback effect of oestradiol, and thus there
The stimulus for the development of these follicles is no LJ-1 surge. At puberty, the threshold is lowered,
is intraovarian and, until the heifer reaches the age thus allowing the pituitary to respond. This is some-
of puberty, they will develop only to the stage where times referred to as the 'gonadostat' theory.
they have a theca interna and then start to undergo Other factors are also involved. The frequency of
atresia. Further development of these follicles to LJ-1 secretion is dependent upon gonadotrophin-
produce mature graafian or antral follicles, of which releasing hormone (CnRH) from the hypothalamus,
there are about 200 growing follicles at puberty which is controlled by an area in the hypothala-
in the heifer, is dependent upon the stimulus of mus referred to as the neural GnRI-I pulse genera-
gonadotrophic hormones (Fig. 1.1 ). Despite the tor. Age-related changes in brain morphology and
absence of oestrous cycles, there is follicular growth, neuronal cytoarchitecture may also be important,
as has been shown using transrectal ultrasonogra- since extrapolation from studies performed in rats,
phy in calves from 2 weeks of age. It was noted that for example, has shown an increase in the number

4
Chapter I 1 I Endogenous and exogenous control of ovarian cyclicity

ofGnRH cells with spine-like processes on the soma the n euroendocrine axis. Although several studies
and dendrites. In addition, the inhibitory effect of h ave shown that leptin treatment can advance the
opioid peptides o n LH secretion is reduced with age, onset o f puberty in both restricted and ad lib fed
which may provide a neurochemical explanation anima ls (Barash ct al 1996, Ahima et al 1997), and
fo r the changes in pituitary sensitivity to oestradiol that serum leptin concentrations increased in pigs
feedback that occur at puberty (Bhanot & Wi lkinson (Qian et al 1999) and heifers (Garcia et a[ 2002),
1983, Wray & Hoffman-Small 1986). it is generally accepted that leptin is not the trigger-
The reason for the 'silent' first oestrus of the ing signal for puberty but a permissive signal that
pubertal animal is believed to be because the cen- enables puberty to occur (Barb & Kraeling 2004 ).
tral nervous system requires to be primed with pro- 'Leptin acts as a metabolic gate; as circulating leptin
gesterone before it will respond and the animal will concentrations increase over the course of pubertal
show be havioural signs of heat. The first ovulatoJy development, serum leptin levels reach a putative
cycle has been shown to be short in pubertal heif- stimulatory threshold which permits activa tion of
ers (7.7 ±0.2 days) and the first corpus luteum (CL) the hypothalamic-pituitary-gonadal axis. This is
not only has a shorter than normal life span but is associa ted with a decrease in the negative feedback
also smaller in size. One explanation for this is that action of oestradiol on the hypothala m ic-pi tuitary
the domi nant follicle, from which the first ovula- axis, and sti mulation of adipose leptin gene exp res-
tion arises, has a lready entered the static phase of sion' (Barb & Kraeling 2004 ).
growth . The su bsequent interovulatory interval is
normal (Adam s 1999). As will be discussed later
Season of the year
in this chapter, o nce puberty has occurred, in most
domestic species there are waves of follicul ar growth In those species that are seasonal breeders, such
and regression. However, even in the prepubertal as the ewe, mare and queen cat, the age at which
animals fo llicular waves have been identified; these puberty occurs will be influenced by the effect of
have been reco rded in heifers (Evans eta[ 1994a, b) season of the year. For instance, a filly born ea rly
and fillies (Nogueira & Ginther 2000). in the year, i.e. January or February, m ay have her
first oestrus in the May or June of the following
year, i.e. when she is 16 or 17 months o ld. A filly
External fadors influencing the foal born late in the year, July o r August, may not
time of onset of puberty have h er first oestrus until she is 21 or 22 months
old. The same is true of ewes, which, depending
The time of o nset of puberty is determined by the u pon the time of year at which they are born, may
individual's genotype, with smaller breeds of animal reach puberty as early as 6 months or as late as 18
tending to be slightly more precocious. However, months o ld.
this inherent timing is influenced b y a number o f
external facto rs. Proximity of the male
Studies in sheep and pigs have shown that exposure
Nutrition
to the male of the species will advance the timing
There is good evide nce that in most domestic spe- of the o nset of puberty. This so-called 'ram or boar
cies, the onset of puberty is closely linked to the effect' is probably mediated by phero mo nal and
attainment of a critical body weight as well as a other sensory cues influencing hypoth ala mic GnRH
minimum percentage body fat. or metabolic mass secretion.
(Frisch 1984 ); thus nutrition is an importanl fac-
tor. Animals that are well fed with good growth
Climate
ra tes reach puberty before those that are poorly fed
with slow growth rates. Hmvever, unl ess the a n i- An thropomorphic extrapolation has assumed that
mal is severely malnourished, cyclical activity will animals living in the tropics reach puberty at an ear-
eventually occur. The protei n lepti n (see below), lier age than those in temperate climates. Studies
secreted by white fat cells in adipose tissue, may ca rried out in Zambia have shown that in cattle this
well provide the link between metabolic status and is not true.

5
 11 I Normal cyclical ovarian activity and its control

Disease Metoestrus
I
Any disease that can influence the growth rate, The phase s~ding oestrus. The granulosa cells of
either directly or because of interference with feed- the ovulated follicle give rise to lutein cells which are
ing and utilization of nutrients, will delay the onset responsible for the formation of the corpus luteum.
of puberty. There is a reduction in the amount of secretion from
the uterine, cervical and vaginal glands.

I The oestrous cycle and its phases Dioestrus

Traditionally, the oestrous cycle is divided into a The period of the corpus luteum. The uterine glands
number of phases. undergo hyperplasia and hypertrophy, the cervix
becomes constricted and the secretions of the gen-
ital tract are scant and sticky; the vaginal mucosa
Pro-oestrus becomes pale. The corpus luteum is fully functional
The phase immediately preceding oestrus. It is during this phase, and is secreting large amounts of
characterized by a marked increase in activity of progesterone.
the reproductive system. There is follicular growth The period of the oestrous cycle when there is a
and regression of the corpus luteum of the previ- functional corpus luteum is sometimes referred to as
ous cycle (in polycyclic species). The uterus enlarges the luteal phase of the cycle, to differentiate it from
ve1y slightly; the endometrium becomes congested the follicular phase. Since in most of our domestic
species oestrus is the only readily identifiable phase
and oedematous and its glands show evidence of
increased secretory activity. The vaginal mucosa of the oestrous cycle, there is some merit, in poly-
oestrous species, in dividing the cycle into oestrus
becomes hyperaemic; the number of cell layers of
the epithelium starts to increase and the superficial and interoestrus, the latter including pro-oestrus,
layers become cornified. The bitch shows external metoestrus and dioestrus. Another alternative divi-
evidence of pro-oestrus with vulval oedema, hyper- sion can be into follicular and luteal phases.
aemia and a sanguineous vulval discharge.
Anoestrus
Oestrus The prolonged period of sexual rest during which the
The period of acceptance of the male. The onset and genital system is mainly quiescent. Follicular devel-
opment is minimal; the corpora lt!tea, although
end of the phase are the only accurately measurable
points in the oestrous cycle and hence are used as identifiable, have regressed and are non-functional.
Secretions are scanty and tenacious, the cervix is
the reference points for determining cycle length.
The animal usually seeks out the male and 'stands' constricted and the vaginal mucosa is pale.
for him to mate her. The uterine, cervical and vaginal
glands secrete increased amounts of mucus; the vag- Natural regulation of cyclical
inal epithelium and endometrium become hyperae-
adivity
mic and congested; the cervix is relaxed.
Ovulation occurs during this phase of the cycle
in all domestic species with the exception of the Regulation of cyclical activity in the female is a com-
cow, where it occurs about 12 hours after the end plex process. With the development of new tech-
of oestrus. Ovulation is a spontaneous process in all niques, particularly those involving hormone assays,
domestic species with the exception of the cat, rab- and the application of new molecular biologi-
bit and camelids, in which it is induced by the act cal techniques, there is a continual advance in the
of coitus. knowledge and understanding of the mechanisms
During pro-oestrus and oestrus, there is follicular involved. Although much of the early work was done
growth in the absence of functional corpora lutea, on laboratory animals - notably the rat and guinea
the main ovarian hormones produced being oestro- pig - there is now much more information about
gens. Pro-oestrus and oestrus are frequently referred domestic species, although there are still areas, par-
to collectively as the follicular phase of the cycle. ticularly in the bitch, that are not fully understood.

6
Chapter I 1 I Endogenous and exogenous control of ovarian cyclicity

The central control of cyclical activity is the hypo- substances. These are secreted by the hypothalamic
thalamic-pituitary-ovarian axis. At one end of this neurons and are carried from the media n emi-
axis there is the influence of the extrahypothalam ic ne nce of the hypothal am us by th e hypothalamic-
areas - the cerebral cortex, thalamus and midbrain - hypophyseal portal system. In 1971 the molecular
and the role played by stimuli such as light. o lfac- structure of porcine GnRH was determin ed {Matsuo
tion and touch (Eilendorff 1978), whi le at the other et a l 1971) as bei ng a decapeptide, and subsequently
end is the influence of the uterus upon the ovary. syn thesized (Geiger et al 1971). Opinion is divided
as to whether GnRJ-1 is responsible in vivo for the
release of both FSH and LH (Lamming et a l 1979),
Melatonin and other pineal peptides
although the injection of GnRH stimulates the
The pineal gland appears to have an important release of both FSH and LH in domestic species. As
role in controlling reproduction in seasonal breed- yet. no specific inhibitory factor such as that for pro-
ing species and also in the timing of puberty by lactin has been identified for gonadotrophins.
influencing the release of FSH, LH and prolac- Specific neurotransmitter substances are involved
tin. Although much of the interest h as been in in the regulation of the release of pituitary hor-
the action of the indo leamine melatonin, there is mones. Thus, noradrenaline (norepinephrine) stim-
increasing interest in the other pineal peptide hor- ulates the release of FSH and LH and the inhibition
mones, namely arginine vasotoci n, gonadotrophin o f the conversion of dopamine to n oradrenaline,
and prolactin-releasing and inhibitory hormo nes. a nd blocks the 'oestradiol-induced' release of LH,
There is some suggestion that melatonin may act which js responsible for ovulation. Serotonin inhib-
not directly upon the hypothalamus/ante rior pitu- its the basal secretion of LH and regulates o ther neu-
itary but indirectly via the other pineal peptide rosecretory systems. Dopamine has an important
hormones. rol e in the control of prolactin release.
Melatonin drives the reproductive response of the There is good evidence that in domestic species
ewe to inductive photoperiods (Bittman et at 1983). the secretion of FSH and LH is controlled oy two
Rhythmic adm inistration of melatonin to adult ewes functionally separate, but superimposable, systems.
exerts a similar effect to increased hours of darkness These are ( 1) the episodic/tonic system, which is
by inducing the onset of the breeding season (Arendt responsible for the continuous basal secretion of
et a l 1983) and causes changes in prolactin con- gonadotrophin and stimulates the growth of both
centrations in the plasma that are similar to those germinal and endocrine compon ents of th e ova1y,
following exposure to sh ort days ( l<ennaway et al (2) the surge system, which controls the short-lived
1983 ). In sheep, an intact pineal gland is required massive secretion of gonadotrophin, particularly
for a normal photoperiodic response to altered day- LH, responsible for ovulation. There are two hypo-
light panerns; however, oth er seasonal environmen- thalamic centres that are involved in con trol ling
tal cues are important, since pinealectomized ewes these two systems (Fig. 1.2).
still show seasonal breeding (Lincoln 1985). With the exception of the cat, rabbit a nd camel ids,
The mare is a seasonal breeder but is 'switched on' all domestic species are spontaneous ovulators.
by increasing day length. The pineal gland is involved, However, in these three species ovulation is induced
since if it is removed the mare does not show a nor- by the stimulation of sensory receptors in the vagina
mal response to changes in photoperiod. In intact and cervix at coitus. This initiates a neuroendocrine
mares, melatonin concentrations increase during reflex ultimately resulting in the activation of GnRH
hours of darkness (Grubaugh et a] 1982). There is neurons in the surge centre and release of a surge
some evidence that foals are conditioned at an early ofLH.
age and develop a pattern of melatonin secretion Not only does the anterior pituitary have a direct
from about 7 weeks of age {Kilmer et al 1982). effect upon ovarian functions by stimulating follicu-
logenesis, follicular maturation, ovulation and cor-
pus luteum formation, but the ovary has an effect
Hypothalamic and anterior pituitary
upon the hypothalamus and anterior pituitary. This
hormones is mediated by oestradiol, produced by the malllr-
The hypothalamus is responsible for the control of ing follicle, and by progesterone, produced by the
release of gonadotrophins from the anterior p itu- corpus luteum. The episodic/tonic hypothalamic
itary by the action ofspecific releasi ng and inhibitory rel ease centre is influenced by the negative- feedback

7
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