SAUNDERS AnimprintofElsevierLunned e EIS(:vit'.rLimih.'l.12001.Allright"reserved. TheriJ::hrofDavtdNoakt'S.l1mllthyParkinsonandGuyEnglandroheidentifiedasauthors{.(this work~l-'beenassertedhytheminaccordancewith theCopvrtghr.Dcsi~,andParent...Act 19M Norartofrhi-, ruhlic:lrion1I1;1)'h:-reproduced.storedinaretrievalsv-tem,tITtransmittedinany formorI:>yilll)'means,electronic,mechanical,phorocopvirur,rccordmaorotherwise,withouteither thepriorpermissionofthepublishersoralicencepermittingrestricted(I'\flying intheUnucd Kingdom i~'o.'llt."l.lt-.ytheC 'l'yrighrLicen-mgA/'.t.'nq,90TonenhamCourt Rll<kl.LoudonWIT-+LP. Pernll~slul~mayhe-~ll.lghldrrectlvfromElsevier'sHealthSciencesRightsDcrenmcmin Phihlt..ldt'hia. USA phone:(+I)2t'52N 1804.fax:(+1)215239 3805.e-m.nl:heahhpt.·rmi""i\lO,,~c1sc\·icr.com. YllUmilY;11"1:1completeY\lllrrequeston-linevia theElsevier hUffiep<tI-'t"(hllr:/I....ww.el-evier.com}, ~yselecnnz'Supponandcontact' aoJ then 'u "l(')Tight andFcrmissioru..•. First{"'l1t>lbhl."\1 1938a-,VeterinaryObsremcshyF.Benesch Secondedition 1951a..VcrcnnarvOh.tetric...hyF. I'\enex-:handJ.G.Wri~ht Thirdedmon 1964a'"Wri~ht'~VerennarvOb-temcsbyG.H.Arthur Fourthedinon 19i; asVercnnarvReproduction andObsrctrtcsbvG. II.Arthur Fifth edmon 1982a..VercrinarvReproductionandOh.tetrics hyG.H.Arthur. D.E.Nuak~anJ H.Pear-on Sixthedmon 1989a..VerennarvReproductionandObstetricshyG. II.Arthur. D.E.NO;lk~andH.Pear-on Reprinted 1992 Seventhedmon1996,h VercnnarvReproducnonandOh.letric.<;t>yG.H.Arthur, D.E.Noakes.H.PearsonandT.J.Parkinson Repnmed1998.1999 Eighthedmon2001a"Arthur'sVcrcnnarvReproductionandObsrctrtcsh~' D.E.Noakes,T.J.Parkin-onandG.C.W.England ReprintedZ003{twice).Z(\..1.+.ZOO;. 2007.2C\.1-8 ISBN,978070102\\6I British Library Catalllj.'.Uinl-:inPublication Data Acataloguerecord(orthishookisavailablefromthe Bnushl.Ihrarv LibraryofConl-:rc!'l~CataloginginPublicationData Acmalo/,.'recordfllrIhil<lhOllkb'I\'ailahlc[rom theLibraryofConJ::tl'~' Note Veterinaryknllwk·d/.:eiscon.,tanrl~'chanJ::ing.Asnew infonn.nionh.·COllWS:I\'aibhlt-,changesin treatment.procedures,equipmentand theuscofdrugsbecomenecessary.Thel'llihlr~/iIlllhor4 contributorsandthepublishershave.asfarasitispossible.taken caretilensurethatthe informationuiveninthistextisaccurateand uptodan-.However,readcrsarel<llroll/.:h·ad\"i~ltu confirmthatthemtorrnanon,espcctallvwith regardtodrugusage.complieswiththelatestlegtslanon andstandard..\lpractice. yoursoun:eforbooks. journalsandmultimedia . . inthehealthsciences www.elsevierhealth.com _. Workingrogerher grow [0 libraries in developingcountries Tho ........""·.cbe\·Kuum I",,,,,,,,,·.boolu.id.org I...""",,·.s.Jh~.org policyIS10use pipetmanufactured fromlustiil'lablttorHtl I PrintedinChina Contributors Dr Nazir Ahmad Professor David E Noakes Associate Professor of Animal Reproduction Professor of Veterinary Obstetrics and Department of Animal Reproduction Diseases of Reproduction University of Agriculture Royal Veterinary College Faisalabad University of London Pakistan North Mymms Hatfield Professor Marzook M Al-Eknah UK Professor of Theriogenology Dr Timothy J Parkinson College of Veterinary Medicine and Animal Resources Associate Professor in Farm Animal Health King Faisal University Institute ofVeterinary,Animal Al-Ahsa and Biomedical Science Kingdom of Saudi Arabia Massey University Palmerson North Mr William B Christie New Zealand Partner Dr Jonathan F Pycock Paragon ET Director,Equine Reproductive Services Tyne Green Messenger Farm Hexham Ryton UK Malton UK Professor Gary C W England Professor of Veterinary Reproduction Mr Martin Sheldon Royal Veterinary College Senior Lecturer North Mymms Royal Veterinary College Hatfield University of London UK North Mymms Hatfield Dr Christianne E Glossop UK Pig Veterinarian Merton Farmhouse Dr Keith C Smith Lea Veterinary Practitioner Malmesbury Department of Animal Health and Husbandry UK University of Bristol Langford Dr Susan E Long UK Senior Lecturer University of Bristol Mr David Whittaker Department of Clinical Veterinary Science Huntingdon Life Sciences Langford Huntingdon UK UK vii Foreword It is with a sense of pride and fulfillment that I have provided welcome encouragement to me and respond to my fellow authors’ to have the eighth fellow authors to continue to update the book. edition of “Veterinary Reproduction and The need to make comprehensive reviews for Obstetrics”entitled to me. student use from the vast volume of new liter- It was in 1962 that the late Professor J.G.Wright ature, particularly in the fields of reproductive invited me to take over his “Benesh and Wright endocrinology, gonadal physiology and genetics, Veterinary Obstetrics”, 2nd edition. In agreeing to is now a heavy task.In the context of stimulus to his request, it was my intention to revise and further effort in authorship is the inspiration from greatly enlarge his text for undergraduate students the work of prominent contributors to one’s par- of veterinary science in English-speaking countries ticular discipline.The period of my 60 year career and for it to serve as a reference source for practis- in clinical veterinary science has been notable for ing veterinary surgeons,as well as for scientists of the contemporary lives of outstanding workers in allied disciplines.These objectives have been fol- my field,from whose publications I have benefited lowed since my 1964 third, to the present eighth, substantially. It is a pleasure to record my deep edition. gratitude to the following gentlemen:- Professor The original format of the book and the divi- W. Williams (Cornell), F. Benesch (Vienna), sional layout of its contents into appropriate sec- N. Lagerlof (Stockholm), J. G.Wright (London), tions and chapters have undergone progressive E.Amoroso (London),M.Vanderplassche (Gent) development but the only major changes in subject and S.J.Roberts (Cornell). titles have been the addition of chapters on “Exotic I will close by expressing my warmest thanks to Species”, “Small Mammals” and “Embryo all who have contributed to previous and present Transfer”,together with a vocational appendix of editions of our book. My special thanks go to the materials used in the diagnosis and treatment Professor David Noakes,who has played a leading of reproductive disorders.The advent of ultrasonic role in his work for the last three editions. imaging has become established as a most useful It is extremely sad to refer to the death of addition to the veterinary surgeons’ armoury; it Professor Harold Pearson since the publication of is appropriately incorporated in the relevant the seventh edition.He is still greatly missed by all chapters of the last three editions. who knew him as a brilliant clinician and teacher. The favourable literary reviews of our book and its wide use overseas,together with translation into Geoffrey H Arthur French,Italian,Spanish,Portuguese and Japanese, 2001 ix Preface It is with great pleasure that we dedicate the 8th chapters devoted specifically to infertility in the edition of Veterinary Reproduction and Obstetrics specific species. We welcome the contributions to Professor Geoffrey Arthur, initially the sole, from four new authors: Dr Nazir Ahmad , and subsequently senior author who successfully Professor Marzook Al-Eknah ,Mr Martin Sheldon husbanded five previous editions of the book from and Dr Keith Smith who revised the chapters on ‘conception to parturition’.It is in recognition of Buffalo, Camels, the caesarian operation in large the outstanding contributions that he has made to animals and infertility in the ewe and goat,respec- the subject over the last 50 years. He has influ- tively.We also thank those contributors to the 7th enced, either directly or indirectly, the profes- Edition,who have painstakingly revised their chap- sional lives of all of the contributors.Although this ters for the 8th edition: Mr Will Christie, embryo will have been primarily in his role as a dedicated transfer; Dr Christianne Glossop, infertility in the and stimulating clinical teacher,it is also because sow;Dr Susan Long,abnormal development of the he was one of the first clinical researchers in vet- conceptus and its consequences,and Dr Jonathon erinary reproduction; this will be recognised by Pycock,infertility in the mare. readers of the book by the frequent reference to In addition, we would also like to express our his published work. thanks to Wendy Lee who did an excellent job as We are sad that the late Professor Harold Pearson Copy Editor,and Sheila Black and Scott Millar of was unable to contribute to this 8thEdition.He was Harcourt Publishers for their hard work and cour- one of the outstanding clinical teachers of the last teous professionalism in producing the 8thEdition 40 years,his writings were based on his vast experi- of the book. ence as a clinician working at the Bristol Veterinary Inevitably, as we complete the revision of the School as shown by his substantial contributions to 8th Edition, thought must be given to a 9th the literature including previous editions of this Edition in the not-to-distant-future.With the vast book. expansion in the knowledge base in the subject, It is intended that this new edition should still and bearing in mind that the principal target cater primarily for the undergraduate, although for the book is the veterinary undergraduate, we others will find it to be a useful reference source.We will need to decide whether we can continue to have added just one additional chapter,and that is enlarge both the length and the scope of the book by Mr David Whittaker who covers normal repro- or whether we must review its format. duction and reproductive disorders in rabbits and rodents,which as well as still having a major role as laboratory animals in research are also being kept David Noakes, in increasing numbers as pets.We have also deleted Tim Parkinson the chapter devoted specifically to Retained Fetal Gary England Membranes and included this material in the 2001 xi Plate 1 Adhesion almost completely obstructing the left uterine horn. Plate 2 Removal of the obstruction by endoscopic cauterisation. (a) (b) (c) (d) (e) (f) Plate 3 Repair of a third degree perineal laceration in a Friesian cow. (a) The perineal laceration exposed with cow under caudal epidural anaesthesia. (b) Commencement of dissection of vaginal mucosa. (c) Completion of dissection of vaginal mucosa; note exposed tissue ready for suturing. (d) Commencement of closure. (e) Closure almost completed. (f) Restoration of a complete shelf of tissue between rectum and vagina, the dorsal commissure of the vulva is subsequently repaired (see Figure 18.5). Plate 4 Placenta of lamb that had aborted due to infection with C. psittaci (ovis). Note cotyledons covered with a light-brown deposit and similar material on the surface of the intercotyledonary chorionic surface. (Courtesy of A. J. Wilsmore). Plate 5 Placenta of lamb that had aborted due to infection with T. gondii. (Courtesy of A. J. Wilsmore). Plate 6 Aborted fetus showing characteristic lesion on the liver of C. fetusinfection. (a) (b) (c) (d) (e) (f) Plate 7 Photomicrographs of exfoliative vagina cells during various stages of the reproductive cycle. The smears have been stained with a modified Wright–Giemsa stain. (a) Anoestrus: parabasal epithelial cells and small intermediate epithelial cells. (b) Pro-oestrus: small intermediate epithelial cells, large intermediate epithelial cells and erythrocytes. Polymorphonuclear leucocytes are also found in low numbers during this stage of the cycle but are not demonstrated here. (c) Early oestrus: large intermediate epithelial cells, anuclear epithelial cells and erythrocytes. Polymorphonuclear leucocytes are generally absent during this stage of the cycle. (d) Oestrus: anuclear epithelial cells, large intermediate epithelial cells and erythrocytes. The percentage of anuclear cells is high. (e) Metoestrus (higher magnification than a–d): small intermediate epithelial cells and large numbers of polymorphonuclear leucocytes. During early metoestrus large intermediate epithelial cells may be present, and later numbers of parabasal epithelial cells increase. There is often a large amount of background debris. (f) Late metoestrus (higher magnification than a–d): parabasal epithelial cells and small vacuolated intermediate epithelial cells are typical of this stage of the cycle but may also be found during anoestrus and more rarely during pro-oestrus. 1 Endogenous and exogenous control of ovarian cyclicity In nature,it is the general rule that animals breed tive activity. Provided the environmental influ- once annually and parturition occurs in the ences are favourable at this time, then once the spring,the time most favourable to the progeny in ‘biological clock’ is started it will continue for as that they grow up during the period of increasing long as the environment remains favourable. In light and warmth,and also at the time when food none of our domestic species is there a physiolo- for the mother is most abundant to ensure ade- gical change comparable with the menopause of quate lactation. Under the conditions of feeding women. and housing provided by domestication the Amongst non-seasonal polycyclic animals,such breeding season tends to be lengthened,and some as the cow and sow,the recurring cyclic activity is of our species, particularly the cattle, may breed interrupted by pregnancy, lactation and patho- at any time during the year; all domesticated logical conditions.In those species which are sea- animals, however, show a constant tendency to sonally polycyclic, the mare, ewe, doe (or nanny) revert to the natural breeding season. goat and cat, or monocyclic like the bitch, there For an animal to breed, it must be mated and are periods of sexual quiescence or anoestrus. hence must attract the male and be sexually When the female reaches puberty the genital receptive (in heat or in oestrus). All domestic organs increase in size. During the prepubertal species show recurring periods of sexual receptiv- period the growth of the genital organs is very ity, or oestrous cycles, which are associated with similar to that of other organ systems, but at the ripening in the ovaries of one or more puberty their growth rate is accelerated, a point Graafian follicles and culminate in the shedding well illustrated in the gilt,where the mean length of one or more ova.If a fertile mating occurs then of the uterine horns is increased by 58%, the pregnancy may ensue. mean weight of the uterus by 72% and the mean weight of the ovaries by 32% between 169 and 186 days of age (Lasley, 1968). Females of PUBERTY AND THE ONSET OF CYCLIC domestic species reach the age of puberty at the ACTIVITY following times: ● mare:1–2 years The young female animal shows no evidence of ● cow:7–18 months recurring or cyclic periods of sexual receptivity. ● ewe:6–15 months The onset of such changes when the female ● doe or nanny goat:4–8 months becomes sexually mature and able to reproduce is ● sow:6–8 months referred to as puberty. Amongst females of the ● bitch:6–20 months domestic species, puberty precedes the develop- ● queen cat:7–12 months ment of physical maturity and, although they become capable of reproducing, their efficiency, The changes that occur at puberty depend particularly with respect to their fecundity, has directly upon the activity of the ovaries,which have not reached its maximum. two functions: the production of the female The initiation of puberty is largely a function of gametes and the synthesis of hormones.Let us con- the animal’s age and maturity since the female is sider the changes that occur in the ovary of the born with a genetic potential for cyclic reproduc- young heifer calf.At birth,each ovary may contain 3
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