Repeat Breeding in Cattle and Buffaloes

CAB Reviews: Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources 2008 3, No.
062
Review
Recent developments in the diagnosis and therapy of repeat breeding cows and buffaloes
G.N. Purohit*
Address: Department of Animal Reproduction, Gynaecology and Obstetrics, College of Veterinary and Animal Science, Bikaner 334001, Rajasthan, India. *Correspondence: Email: gnpvog@yahoo.co.in
Received: Accepted: 15 May 2008 15 July 2008
doi: 10.1079/PAVSNNR20083062 The electronic version of this article is the denitive one. It is located here: http://www.cababstractsplus.org/cabreviews
g
CAB International 2008 (Online ISSN 1749-8848)
Abstract Repeat breeding (RB) continues to be a big problem for breeders and veterinary clinicians. A brief mention is made of the common aetiological and risk factors for RB in cattle and buffaloes, and the possible diagnostic and therapeutic approaches are described in detail. Important diagnostic tools could include rectogenital palpation, vaginoscopy, uterine cytology and the in vivo imaging technique of ultrasonography. When considering the most common causes of RB, vaginoscopy and palpation continue to be the only diagnostic tools available to clinicians at many locations, while by using ultrasonography, diagnostic accuracy can be increased markedly, especially when dealing with individual cows or buffaloes. Contrarily, when dealing with herds, metabolic proles and sampling to detect infectious disease must be the clinicians choice. Of pertinent consideration are the management regimens and feeding practices. Despite the development of many diagnostic procedures such as hormone assays, colour Doppler sonography, and hysteroscopy, diagnosing the cause of pregnancy failure in an individual cow/buffalo continues to be difcult, as a proportion of animals demonstrate obscure infertility. The choice of a therapeutic regimen depends on the possible cause of RB. Recent advances in the therapy of endometritis include the use of immunomodulators such as Escherichia coli lipopolysaccharide, use of eicosanoid PGF2a and therapy with enzymes with or without therapy with antibiotics, the use of which continues to be debatable. The therapy of ovulation induction in various ovulatory disturbances includes regimens utilizing hCG, GnRH, prostaglandins and their combinations. It appears that RB animals with aberrations of oestrus cycle do demonstrate such ovulation asynchronies. Suprabasal progesterone concentration at oestrus is thought to be an important contributor of RB, but remediation of this is largely unknown although reducing stress appears to be a probable method. Luteal insufciency can be resolved by administration of hCG and GnRH or progestagens. A brief mention is made of ways of improving management and insemination procedures. Keywords: Cow, Buffalo, Repeat breeding, Diagnosis, Treatment
Review Methodology: We searched the following databases: CAB Abstracts, Animal Breeding Abstracts, and PubMed (keyword search terms used: repeat breeding, in vivo imaging, immunoinfertility, endometritis, ovarian cysts, infertility, endoscopy and metabolic proles). In addition, we used the references from the articles obtained by this method to check for additional relevant material. We also spoke to colleagues and checked for any upcoming studies not yet published.
Introduction The repeat breeding (RB) syndrome continues to be a major problem in cattle and buffalo breeding, leading to large economic losses to the dairy producers [1, 2]. Some authors consider RB to be overemphasized and that
modern high-producing Holstein cows have reduced fertility because of intensive selection for high yields [37]; however, others do not concur with this view [8, 9]. Recently, RB cows have been dened as a heterogeneous group of subfertile cows with no anatomical abnormalities or infections that exhibit a variety of reproductive
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Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources
disturbances in a consistent pattern over three or more consecutive heat cycles of normal duration (1725 days) [10]. Concurrence does exist for a similar denition in the water buffalo [11], as cattle and buffalo are an integral part of the mixed-crop-livestock smallholder farming systems in the developing countries of the Asia-Pacic region [12]. However, buffaloes are considered tangentially seasonal animals and their reproductive efciency is usually negatively affected by increasing the length of daylight, with the obvious exception of the equatorial regions, where the reproductive function is mostly dictated by the availability of feedstuff rather then length of light hours [1317]. Because of these and other subtle differences between cattle and buffalo (fertility in buffaloes is considered lower than in cattle), [18] essentially therefore RB in the buffalo must be considered only during the breeding season. It has long been argued that the causes of the RB syndrome are either failure of fertilization [19] or early embryonic deaths [2023], with embryonic deaths accounting for the major share of the reproductive wastage in dairy cattle [21, 2434]. However, such descriptions are few for the buffalo [35] but the phenomena are usually considered to be similarly existent [11, 36]. More recently, it has been considered that the cow, the bull and a range of environmental and handling factors often overlapping each other result in RB and it is often difcult to determine the primary origin [10]. Such problems are much more difcult to trace when a farmers individual cows or buffaloes from a variety of management practices have to be investigated instead of investigations on a herd. In general, the pregnancy rates are higher where natural service is the method of breeding compared with articial insemination [37, 38]. Therefore, RB has also to be viewed in this perspective. In general, the veterinary clinicians in many developing countries face the problem of treating such problem cattle and buffaloes with little to diagnose and lack of a systematic therapeutic regimen. This review focuses on the possible diagnostic modalities and therapeutic regimens in RB cows and buffaloes.
approaches, a brief mention is made of the possible aetiological and risk factors. The aetiology of RB appears to be multifactorial. All or any of the causes described for pregnancy failures in cattle and buffaloes are evidenced clinically in the form of RB. When evaluating females for RB, the causes for fertilization failures in part or in toto that would result from male gamete abnormality or hypofunction must be rst eliminated. The possibility of failures caused by semen abnormality appears to be ruled out when articial insemination using semen of high fertility is used. However, it must be kept in mind that frozen semen has a shorter life span, < 12 h in the female tract, compared with ejaculated semen [45] and a lower fertility compared with fresh semen because of lower viability post thaw and sublethal dysfunction in a proportion of the surviving subpopulation [46]. A problem with buffalo semen appears to be the season during which semen is collected, because semen collected during hot summer months appears to offer suboptimal fertility compared with that collected during the winter months [4750]. Such seasonal variations, however, are not known to cause deleterious changes in sperm quality in swamp Thai buffaloes [51]. A brief mention is made herewith of the possible aetiological factors that can contribute to the RB syndrome in cattle and buffaloes.
Nutritional Inadequacies Some of the nutritional deciencies that are known to result in pregnancy failures include lack of energy [5260], excess of dietary protein [6163], and deciencies in micronutrients [64] such as calcium, phosphorus and iodine [6569], cobalt, copper, zinc and magnesium [65, 70], vitamin A [7174] and selenium and vitamin E [69, 7577]. Vitamin A and b-carotene [3, 78] deciency or excess of body metabolites such as glucose, urea, albumin, globulin, and non-esteried fatty acids may directly or indirectly affect follicle growth, conception and embryonic development. In buffaloes, only a few descriptions [79, 80] of inadequate nutrition and RB are available; moreover, the important aspects of nutritional management of dairy cows for optimum postpartum fertility [8184] appear different from those found in buffaloes, as more important for cows is the resumption of postpartum oestrus, which occurs 90 days after calving only in 3949% of buffaloes, the rest remaining in anoestrus for 150 days [16].
Incidence For various countries, the incidence of the problem has been described in cattle and buffaloes to range from 5 to 35% [21, 3941]. However, such an assessment of incidence suffers from limited data over a few or many herds. A seasonal inuence on the appearance of the problem has been depicted, with the hot season being unfavourable. The incidence of RB in cattle and buffalo in various reports is presented in Table 1.
Hormonal Dysfunction The entire events of pregnancy establishment from ovulation of a viable competent oocyte to fertilization, to implantation and subsequent growth of an embryo in utero are dependent on a complex chain of rhythmic hormone secretion and binding [85]. A slight deviation in any of
Aetiology and Risk Factors The aetiology of RB has been widely reviewed [1, 22, 23, 4244] but to correlate diagnostic and therapeutic
G.N. Purohit
Table 1 Incidence of repeat breeding in cows and buffaloes in various studies Incidence (%) Cows 14.427.0 10 5.533.3 1015 21.428.2 18.124.4 16.5 16.418.8 5.0 16.658.8 8.98 8.98 6.8 19.8 8.209.30 725.0 7.418.6 25.9 5.0 24.0 8.0 3.0 15.8 8.33 10 4.2 9.77 17.8 10 7.3 25.1 28.4 717.0 12.0 Buffaloes 7.37 10.76 6.0 520% 8.0628.84 8.33 1.9 6.1 7.57 Breed Hariana Swedish breed Red Sindhi HF Red Sindhi Sahiwal Tharparkar Crossbred HF Fulani Cows JerseyHF Crossbred Crossbred JerseyGir HFTharparkar Cows, buffaloes Danish Red, Sahiwal, HF HFGir HF HF Red Kandhari Crossbred Crossbred Egyptian cows HF HFDeoni Crossbred Crossbred HF Crossbred HF SahiwalFriesian Rathi HF crossbred Jaffarabadi Murrah, Nilli-Ravi ND, Murrah Murrah ND, Murrah, Nilli-Ravi Egyptian ND, Murrah Mixed Mixed Type of management FM FM FM FM FM FM FM FM FM VM VM VM VM FM FM FM FM FM FM FM FM FM FM FM FM FM FM FM FM VM FM FM VM Clinics FM FM VM FM Clinics VM VM VM VM Reference [469] [42] [470] [178] [471] [471] [471] [471] [472] [473] [474] [41] [475] [476] [477] [478] [190] [479] [480] [1] [481] [481] [482] [3] [28] [483] [484] [485] [29] [486] [487] [488] [280] [489] [490] [40] [41] [491] [492, 493] [494] [486] [37] [495]
HF, Holstein Friesian; FM, farm management; VM, village management; ND, Non-descript.
the hormones may change or affect the establishment of pregnancy. Disturbances of ovulation in part because of faulty luteinizing hormone (LH) secretion, a prolonged duration of standing oestrus or improper steroid genesis appear to be major causes of RB in dairy cattle [8688]. An important feature of buffalo reproduction appears to be the levels of plasma prolactin, which are elevated during summer and known to suppress progesterone concentrations [7, 89] besides low luteal progesterone that accounts for 50% of embryonic mortalities in buffaloes during summer [35]. Besides the importance of optimal progesterone concentrations during the luteal phase, which may depress thyroid function during summer
[90, 91] and culminate into pregnancy failures because of early embryonic deaths or lack of signal transduction between the mother and embryo, importance is currently attached to the suprabasal concentrations of progesterone (higher basal progesterone at oestrus) at oestrus in dairy cows [9298]. Such reports are largely unavailable for the buffalo. Moreover, in the buffalo the steroid secretion is inherently low at oestrus [99] and behavioural oestrus is associated with high progesterone levels [100, 101]. A higher progesterone results in poor conception rates largely because of ovulationinsemination mismatch. Ovulatory disturbances commonly encountered in RB animals include delayed ovulation [86, 102] anovulation
[88, 103105] and ovarian cysts [106110]. A manifestation of hormonal dysfunction could be aberrant oestrus expression in dairy cows [6, 10].
Infectious Agents and Reproductive Tract Abnormalities Various reproductive tract abnormalities have been described to be a reason for RB both in cattle [20, 111114] and buffalo [115, 116]. Only a few of these described conditions can be discerned clinically, such as infections of the tubular tract from vagina to fallopian tubes or ovarian cysts and sometimes ovarobursal adhesions. Infectious agents present in the genital tract may hamper fertilization and early embryo development as well. These organisms when present in pathological numbers may produce some toxins or render the uterine milieu unfavourable for conception. Infections must be suspected when there is a moderate to mild degree of endometritis, evident because of akes of pus or occulent material in cervico-vaginal mucus discharge at oestrus. However, when the infections are subclinical they remain obscure clinically but still may hamper conception. A wide variety of bacteria, viruses, fungi and protozoa have been revealed to hamper conception and have been widely reviewed elsewhere [30, 117123], with bacteria being a particularly common problem.
stressors, e.g. high milk production, postpartum disorders and negative energy balance, inammations and infections, lameness, social factors, transport and heat stress. High milk yield, high parity and calving in winter were risk factors for several reproductive disorders, which in turn delayed insemination and conception in dairy cows [155].
Diagnostic Methods In view of the wide variety of causes that can result in RB, the diagnostic procedures in the present review have been classied into the following groups: 1. 2. 3. 4. 5. Record analysis Visual Recto-genital palpation Vaginoscopy Tests to evaluate uterine health: (a) Uterine pH (b) Uterine microbiology (c) Uterine biopsy and cytology Metabolic proles In vivo imaging techniques Immunological tests Endoscopy Tubal patency testing Hormone assays
6. 7. 8. 9. 10. 11.
Genetic Problems and Immunoinfertility Record Analysis Chromosomal abnormalities are considered by a few authors as aetiological agents contributing to conception failures [24, 124128]. Immunological incompatibility of the sperm and oocyte because of production of anti-sperm antibodies has been documented [129142] in both cattle and buffaloes to be one reason for fertilization failures. Analysis of records when traced retrospectively would provide the number of actual inseminations and previous periparturient disease that have resulted in suboptimal fertility. Insights into poor fertility can be traced in individual cows by record analysis. However, more often, cows/buffaloes presented to clinicians with RB originate from diverse changing management strategies, with no records.
Miscellaneous High environmental temperatures [143, 144], season, size of herd/type of housing, age [143148], environmental pollutants [149], milk yield, lactation and difcult calvings [21, 150, 151], metabolic disorders [81], postpartum metritis and ovarian cysts [152] are a few of the other risk factors that may increase the incidence of RB in cattle and buffaloes. Stress has been addressed as a cause of impaired reproductive efciency [153] and the hormonal mechanism for effect on fertility is common irrespective of the stressor involved. In a stressful situation, the function of the hypothalamuspituitarygonadal axis might be disrupted at each level [154]. Many factors in modern dairy farming have been identied as potential Visual The importance of visual observations in diagnosing an animal that would subsequently repeat to service or articial insemination (AI) lies in the fact that human factors such as improper oestrus detection or insemination asynchrony may many times contribute to the failure of pregnancy establishment in an individual animal. Cruz [156] has stressed that the poor results in insemination programmes have been largely the result of human factors such as improper insemination techniques or improper timing of AI. Visual observations that need the attention of the inseminators include the colour, consistency and
G.N. Purohit
clarity of the cervico-vaginal mucus, vulvar oedema and vaginal congestion. A cow showing a cervico-vaginal mucus discharge that is watery, thin or copious should be viewed as one with suspicious subsequent fertility. In natural mating programmes, which are more readily practised in the buffalo, size differences in the male and female or services without any ejaculatory thrusts may result in conception failures. The incidence of RB with individual bulls has been commented on recently [157], and it must be kept in mind that when more females are failing to conceive to a particular male, the reason often lies in that male which must be investigated and it is often better to replace it. An uncommon condition that is often missed by insemination personnel is uro-vagina. The urine pool of vagina contaminates the semen, often reducing its fertility. A thorough vaginoscopic examination must be undertaken in animals that have doubtful fertility to exclude the possibility of any vaginal growths, scars or adhesions that may impair fertility.
A common belief that pregnancy diagnosis by rectal palpation between days 3541 by the foetal membrane slip can often result in foetal death has been proved to be wrong when palpations are preformed with care [163165]. Rectal palpation can help in diagnosing grossly enlarged fallopian tubes, but enlargements and constrictions of minor nature cannot be evaluated by rectal palpation alone. Likewise, ovarobursal adhesions can be diagnosed by recto-genital palpation [22, 23].
Vaginoscopy Examining the vagina and cervix by vaginoscopy to determine the presence or obsence of small quantities of infected material inside has been a common clinical method to estimate uterine infection. Miller et al. [542] concluded that vaginoscopic examination is a more accurate method for detecting uterine infections than palpation per rectum. Cows with abnormal vaginal discharge on vaginoscopic examination have poor reproductive performance [166, 167]. However, vaginoscopy often fails to identify all cows that are truly at risk of poor reproductive performance and the absence of discharge at vaginoscopy does not necessarily indicate absence of uterine inammation [168, 169]. The presence of discharge in the vagina and its identication by vaginoscopy may be inuenced by the severity of the infection, myometrial contraction, uterine clearance mechanisms, perineal conformation, body condition, postural changes and exercise. Discharges may not be detected in cows in which the cervix is closed, although these cows may harbour infection. A single vaginoscopic examination therefore lacks accuracy and may result in undiagnosed and untreated endometritis [168]. An alternative approach for sampling of vaginal contents using a novel device termed Metricheck was found to be more sensitive in detecting endometritis compared with vaginoscopy [170].
Recto-genital Palpation Recto-genital palpation of both uterine horns, cervical os and the ovaries is by far the commonest diagnostic method used for cows and buffaloes repeating to natural services/inseminations, yielding little information as to the cause of pregnancy failures. However, a systematic approach would denitely give some clue to this multi factorial problem. Animals with poor uterine tone at insemination often have poor conception; hence this must receive attention but not guiding. Moreover, such an evaluation is subjective and often graded differently by different clinicians. Pleuriparous cows, especially those which have had some periparturient problem, have irregularly shaped cervices, creating difculty in the introduction of the insemination pipettes. Such problems appear to be rare in buffaloes. The optimum size of the ovulatory follicles at AI has been commented on elsewhere [158]. However, such evaluations are far from perfect by rectal palpation, and extremely difcult for the buffalo. An important event during pregnancy establishment is the ovulation of the follicle. For estimation of ovulation, palpations must be done every 12 h from AI till the nding of an ovulation depression. Such an evaluation again is difcult for the buffalo. Evaluating the early or late corpus luteum (CL) to rule out luteal insufciency is subjective and the predictions are often suboptimal. Prediction of early foetal deaths by rectal palpation when it occurs beyond day 45 gestation is often possible, but then such deaths often result in voiding of foetal uids, blood and/or foetus itself and is considered an abortion rather than an early embryonic death. Moreover, much of the loss of potential offspring in cattle is concentrated during the rst 42 days after breeding [32] and in particular 620 days after breeding [24, 159162], when it is seldom possible by rectal palpation to evaluate pregnancy.
Tests to Evaluate Uterine Health Uterine pH The pH of the uterine lumen during different stages of the oestrus cycle varies widely, with the lowest pH occurring 2 days prior to ovulation; however, at oestrus it is known to be 7.30 [171]. Samples of uterine secretions are difcult to collect although a few studies point out that this can serve as a partial indicator of the uterine milieu where normal gamete transport and development of the embryo can occur. A reduction in pH from 7.2 to 6.97.1 has been reported to compromise embryonic development [172, 173]. In contrast, cows/buffaloes with metritis show a pH
from 8.23 to 8.80 [174, 175]. The pH of vaginal mucus was 8.5+1.16 in repeat breeder cows compared with 7.2+1.10 in normal breeding cows [132]. An important aw in the estimation of the uterine pH is the lack of development of special sensing probes that can directly be placed in the bovine/bubaline uterus. Feeding diets high in protein alters the uterine environment by reducing concentrations of magnesium, potassium and phosphorus in uterine secretions [61] and by reducing uterine pH [62, 176].
time-consuming and often inconclusive in diagnosing the cause of RB in an individual animal.
Uterine biopsy and cytology Carefully performed uterine biopsies can often reveal the changes in the endometrium and the extent of cellular inltration and/or cellular morphology changes [185]. However, they can seldom help in formulating therapeutic measures and therefore their use is, and must be, reserved for forbidden cases suspected for uterine growths, enlargements or malfunctions, in which they supplement as substantial evidence for removal/discard of such animals. In RB cows the glandular secretions and supranuclear vacuolations are observed in histological sections prepared from collected biopsies [186]. Changes observed in cows with endometritis include denudation of epithelial lining and inltration of lymphocyte and neutrophils [177, 187189]. Biopsies of RB buffaloes have also revealed endometritis of varying degree [190]. The endometrial EGF concentration is altered in RB cows and can serve as a potential marker for the identication of cows that would turn out to be repeat breeder [191]. The sensitivity and specicity of uterine biopsy for pregnancy was found to be 92 and 77% [188]. The histological ndings of inammatory changes and brosis were correlated with presence of bacteria [192194]. There is a growing body of evidence on the use of uterine cytology as a means of evaluating uterine health [168, 169, 195, 196]. There is an increase in the percentage of polymorphonuclear leucocytes (PMNs) during clinical and subclinical uterine inammation. Different procedures have been described for obtaining the uterine cells and performing a count, and include ushing the uterus with small amounts (25 ml) of uid [169] or using a commercially available cytobrush [197]. When using a uid recovery procedure after infusion into the uterus, the uid must be centrifuged to concentrate the cells in a small amount of medium in order to enable effective cellular concentration and a good slide preparation. There appears to be lack of standard for identication of uterine health and labelling of endometritis based on endometrial cytology. The cell counts vary with respect to days postpartum. The threshold for dening subclinical endometritis was nding of PMNs >18% at 2033 days postpartum, whereas the respective threshold at 3947 days postpartum was >10% [168]. Between 40 and 60 days of parturition the threshold for endometritis was only 5% [169]. RB cows and buffaloes usually presented to the clinician belong to a diverse group of animals that had calved from 60 to 120 days previously. It therefore remains to be seen how endometrial cytology can help in dening subclinical endometritis in these animals. Moreover, a simple technique that can be used by most clinicians would require simplicity of technique and specicity and consistency of interpretations.
Uterine Microbiology A wide variety of microbes normally harbour the uterine lumen and only when their numbers are high are they evident clinically in the form of purulent mucus akes of pus and changes in odour of the genital discharges. It has been shown in many studies that mostly mixed infections are present in the uteri of RB cows and buffaloes [177 180]. In the authors experience, a subclinical uterine infection remains clinically obscure as also observed in some previous studies [181183] and this appears to be one of the leading causes of fertilization failure both in cattle and buffaloes. Diagnostic tests to evaluate such subclinical uterine infections (endometritis) have been developed to a limited extent, but suffer from incorporation of suboptimal uterine secretion collection methods and accuracy of diagnosis. One such test, the white side test, uses cervical mucus of suspected cows (with metritis/endometritis), which is heated with sodium hydroxide solution up to boiling point. The reaction is considered positive if the colour turns yellow. A correlation between the number of leucocytes present in the mucus and the intensity of yellow colour is the basis of this test [175, 184]. Other recent developments to evaluate endometritis include the novel intravaginal device Metricheck, mentioned earlier, which is known to be more sensitive in detecting endometritis than vaginoscopy [170]. The device consists of a 40 mm hemisphere of silicon attached to a 500 mm long stainless steel rod. The device is inserted through cleaned vulvar lips, advanced to the cranial extent of the vaginal fornix and then retracted back. Purulent material may be visualized within the concave surface, or adherent to the convex surface, of the device. Periparturient problems often contribute to some microbes being harboured in the genital tract, resulting in deviant fertility of obscure nature. Microbes such as Campylobacter are difcult to isolate, but their relative presence is insignicant with the widespread use of AI. When fertility of a herd is in question, it is often desirable to collect random samples from the uteri of some cows using proper techniques in order to nd out the probable aetiological agent. However, such tests are
G.N. Purohit
Table 2 Blood biochemical constituents in normal breeding and repeat breeding cows and buffaloes in various studies Cows Parameter Glucose (mg%) Cholesterol (mg%) Hb (gm%) Ca (mg%) P (mg%) Fe (mg/dl) Mn (mg/ml) Zn (mg/ml) Cu (mg/ml) Blood urea (mg%) Vitamin A (mg/dl) Co (mg/ml) Na (meq/l) K (meq/l) Cl (meq/l) Mg (mg%) Normal 47.1684.54 83.0249.22 9.0611.74 6.1710.73 4.228.19 1.92.4 0.460.58 1.093.14 0.651.14 18.80 41.216 2.189.67 133.7 4.44 96.0 3.1922.57 RB 45.697.73 77.8182.37 8.989.71 6.6069.65 3.378.03 2.4711.3 0.170.19 0.651.19 0.220.99 28.88 37.14 0.85 140 4.27 96.1 2.569.82 Reference [398, 525527] [527529] [527, 530, 531] [529, 532534] [525, 532, 533] [532, 535, 536] [526, 532] [526, 531, 534] [531, 534] [526] [532] [534] [536] [536] [536] [533, 534] Buffaloes Normal 62.590.00 40.23144.98 7.89.4 9.3315.00 4.58.99 0.030.62 0.171.00 0.140.88 0.02 182184 6.537.4 3.303.68 RB 52.582.5 52.1573.01 78.6 916 5.58.0 0.88 0.62 0.16 Reference [537] [537, [537] [537, [537, [539] [539, [539, [541] [539] [539] [539] 538] 539] 540] 541] 541]
Metabolic Prole There appears to be a complex mechanism involved in the interplay of various serum metabolites: the macro (glucose, total protein and lipids) and micro (calcium, phosphorus, various vitamins and trace minerals) nutrients with the different reproductive events; however, because there is an indirect neuro-hormonal mechanism [52], it is difcult to establish concrete clinical norms that would predict potential fertility. The importance of negative energy balance on reproduction has been stressed elsewhere [59] as has been the impact of heat stress [143]. An important parameter that could have some diagnostic signicance could be circulating blood glucose, as low levels are known to affect oestrogen production by the dominant preovulatory follicle [59] and levels of IGF-1. Table 2 depicts the serum biochemical metabolites reported for normal breeding and RB cattle and buffalo in various studies. In one study, the blood metabolites globulin, albumin, urea and b-hydroxybutyrate did not correlate with reproductive performance in dual purpose cows in Mexico [198]. However, this is not always true with RB cattle. A metabolic prole analysis of various blood parameters, such as blood haematocrit, glucose, cholesterol and calcium, can diagnose malnutrition and therefore be useful in high-producing dairy cows [199]. The author partially concurs with this view and suggests a complete blood biochemistry to be undertaken in RB animals wherever possible.
In Vivo Imaging Techniques By far, the most important diagnostic modality for reproductive diagnosis is ultrasonography (USG). The diagnostic signicance of USG lies in the fact that the technique is non-invasive, and ovarian and uterine
morphological changes otherwise undetected by techniques like rectal palpation can be detected and traced [200]. Follicular growth pattern in RB cows revealed that such cows more frequently had two follicular waves corresponding to longer cycles [201]. The CL becomes visible by USG after 3 days of ovulation [202], and USG is considered reliable for measuring follicles and detecting CL [203]. Moreover, the health of the uterus can be evaluated. At oestrus, there is distinct folding of endometrium, and in uterine inammation, echogenic snowy patches are visible sonographically [204]. Ovarian dysfunction is known to be common in RB animals [10] and includes ovarian cysts, ovulation defects, luteal dysfunction and a prolonged life span of pre-ovulatory follicle [98]. Ovulation can be traced by regular scanning at least at 12 h intervals from AI. This would rule out ovulationinsemination asynchrony. Cows and buffaloes not ovulating within 24 h of an insemination must either be re-inseminated or considered for an ovulation induction treatment along with AI. A single USG examination, however, cannot detect ovarian function, and therefore repeated examinations are necessary. In RB cows the USG examination therefore must be done at AI to determine the presence of an ovulatory follicle and then repeated at 12 h intervals to nd out if ovulation has occurred, and subsequent examinations have to be done at 4 day intervals to observe CL formation [10]. Although variable, the optimum size diametric of a follicle at AI would be 1.52.0 cm in dairy cows and 0.91.8 cm in buffalo; however, it has been shown that follicle size has no effect on fertility when ovulation occurs spontaneously [158]. USG can delineate cows/buffaloes with subclinical uterine infection (endometritis). A uterine lumen with a diameter of 0.2 cm and presence of echogenic content in the uterus is considered to indicate endometritis and is known to have a signicant negative association with
conception rate and proportion of cows pregnant [168, 205]. Other important determinants of pregnancy failures that can be detected by USG are luteal formation and embryonic deaths. The morphological characters of CL on both the bovine and bubaline ovaries can well be visualized by day 5 of oestrus. However, this does not precisely predict functionality of this temporary endocrine gland. It is known that pregnancy can be diagnosed with around 100% accuracy at day 25 [206] and the embryonic heart beat can rst be visualized at around day 25 of gestation [207]. It is therefore possible to trace death/ resorption of embryos, which is otherwise not possible by any other diagnostic modality. The maximum early embryonic deaths in cattle occur by day 20 [161]. Late embryonic deaths (day 2742) account for 1020% of embryonic deaths in cattle [208], and these could be more readily detected by USG. The presence of amniotic vesicle/foetus at a particular moment by USG and disappearance at a later time clearly suggests embryonic death. Improvements in the in vivo imaging technique [209] include computer-assisted image analysis of USG [210 214], three-dimensional USG [215], colour Doppler USG [216221] and magnetic resonance imaging (MRI) [222 227]. A better understanding of the picture components obtained by USG can be obtained by image analysis by computers. The data obtained from sonographic assessment are standardized and incorporated into computer analysis software. These modalities widen the understanding of the sonograms. Colour Doppler USG is meant to demonstrate changes that occur in circulation to the uterus, ovaries or ovarian structure and hence can provide new information about physiological changes that occur in the genital organ. The application of these and other in vivo imaging techniques such as MRI has widened our understanding of basic reproductive processes. Prototypes of MRI instruments for intravaginal or intrarectal use are being developed to make this technique more user-friendly [209]. However, due to the high cost of these equipments and the skills required, the use of these modern in vivo imaging techniques is currently limited and is beyond the scope of diagnosis in RB animals.
hormone needs repeated blood sampling and costlier methodology (such as radioimmunoassay (RIA), [229]) that makes such assays impractical under most bovine and bubaline practice. The levels of progesterone both at oestrus and during the luteal phase appear to be critical from most studies in dairy cattle [30, 34, 230232] and buffaloes [35, 233]. Higher basal progesterone (the socalled suprabasal (SB) progesterone) can be considered as a tool for the identication of repeat breeder heifers [95, 96, 234, 235] and buffaloes [93], provided that heat detection and AI timing are optimal. It has been shown that as the progesterone level at AI rises, conception rate in cows declines [236]. The season appears to have distinct effects on buffalo endocrinology, especially the thyroid and prolactin secretions. The thyroid function appears to be depressed during summer and in poorly reproducing buffaloes [91]. Buffaloes with low-plasma protein-bound iodine had low progesterone and a higher number of services per conception [90]. Similarly, prolactin and progesterone are negatively correlated during summer [7, 89]. High levels of prolactin during summer and low LH result in poor reproductive efciency in buffaloes during summer. Hormonal proles in the course of oestrus cycle are on the whole similar in cattle and buffalo [17]. In spite of the impact of seasonal inuences on buffaloes, progesterone production appears to be crucial for embryo viability in buffaloes [35], similar to cattle, and both a late post-ovulatory progesterone rise and low luteal phase concentration are associated with poor embryo development and production of insufcient interferon IFN to preserve luteal regression [30, 35, 102, 237241]. It can probably be concluded from a large number of published studies that whenever possible, progesterone assays must be done on individual cows/ buffaloes at AI and 7 days later. They can serve as diagnostic parameters as to whether the animal would conceive and continue the pregnancy or not. Such inferences, however, need to be validated experimentally with larger trials.
Immunological and In Vitro Tests Immunoinfertility appears to have received more attention in human species. Antisperm antibodies are known to be present in serum of both sexes in human studies [242 245] and they result in sperm-immobilizing activity, leading to penetration reduction of sperm in cervical mucus and resultant reduced fertility [245247]. The detection of these antibodies in cattle is usually carried out by determination of these antisperm antibodies in serum [137, 146, 147] or cervical mucus [129, 130, 132, 134] and similarly in buffaloes [135, 138, 140]. Farahani et. al. [131], however, found agglutinating (Aggl) and immunouorescent (IF) antibodies in serum from repeat breeder, fertile cows and virgin heifers with no
Circulating Hormone Assays Perturbations of the reproductive hormones, especially LH and the ovarian steroids oestrogen and progesterone, can affect pregnancy establishment. A delay/deviation in the secretion of LH peak surge can affect ovulation and the development of CL, but such deviant secretions appear to be multifactorial (e.g. linked to stress, heat, insufcient oestrogen by the follicle, and high prolactin). In buffaloes, a consecutive LH peak, usually accompanied by a double ovulation, has been recorded to be occurring in a small proportion of animals [228]. Clinical assays of this
G.N. Purohit
sperm-immobilizing antibodies. The antibodies were assumed to be produced naturally with no need of female exposure to sperm antigens as all virgin heifers also demonstrated Aggl and IF antibodies in their serum. Their study pointed out that antibodies against sperm are not responsible for reduced fertility in RB cows. Contrary to earlier reports in buffaloes, Kanchev et al. [138] also found that circulating agglutinating antisperm antibodies are very rarely detected in the buffalo cows with unexplained infertility after several inseminations. The role of antisperm antibodies therefore in RB cows/buffaloes remains suspicious and hence puts a question mark on such tests. Methods of predicting the in vivo potential fertility of frozen bull/buffalo bull semen could include the cervical mucus penetration assay [248, 249] or penetration of sperm in polyacrylamide gel [139]. These tests and tests like those performed on oocytes such as the zona binding assay and hemizona assay [250, 251] and penetration of zona-free hamster oocytes by bull sperm [252] in combination can provide valuable information about a semen donor, an insemination dose or a method for semen preservation [253]. However, a pregnancy remains the most appropriate test of potential bull fertility.
Endoscopy A potential means of evaluating morphology and functional means of live tissues could be the endoscopic visualization of the uterus and other genital organs. Endoscopy has been used to a limited extent for visualization and surgical intervention in cows and buffaloes. The use of a exible bre-optic endoscope for clinical assessment of the uterus and intrauterine therapy has been described for the mare [262]. However, similar reports are very few for cattle [263]. Direct hysteroscopy has been used to evaluate the uterus. A paediatric gastroscope (130 cm9.5 cm) and air insufation allow good visibility of both uterine cornua [264]. The only difculty experienced in cows is traversing the cervical canal [264], which has in fact prevented the widespread use of hysteroscopy in bovine reproductive diagnosis. Endoscopic techniques using a ank approach and a 60 cm10 mm rigid endoscope (laparoscope) have been used to view the ovaries of conscious cattle [265, 266] and buffaloes [267, 268] employing CO2 insufation and a head-down tilt. In addition, endoscopy has been used to examine the ovaries by colpotomy [269] or ank methods, particularly for follicular aspiration [270272] or oviductal transfer of in vitro produced embryos [273]. The general principles of laparoscopy [274, 275] and laparoscopic surgeries in adult cattle have been described recently [276]. Laparoscopic ovariectomy in standing cows has similarly been described recently with indications for use in research and removal of tumour-affected ovaries [277].
Tubal Patency Testing The fallopian tube is a particularly complex structure and, as such, an ideal method for its clinical assessment is very difcult to obtain [254]. The incidence of fallopian tube lesions is known to be 6.85% (range 2.69.0%) with ovarobursal adhesions being the commonest lesion in cows [255]. The occlusion of the fallopian tubes can result in lowered fertility when it is unilateral and sterility when it is bilateral [22, 23]. The occlusions may not essentially be because of any lesion inside [256]. An instrument for diagnosing oviduct patency in cows was developed a long time ago [257], with many subsequent modications. The usual test utilizes infusion of phenol sulpfopthalein (PSP) dye using a two-way catheter into the uterine horn and detection of the dye in urine. In animals with nonoccluded fallopian tube, the dye is present in urine within 30 min; however, in cows with non-patent (occluded) fallopian tube, the dye is not visible in urine for up to 2 h [258260]. When the same procedure is to be repeated in the other horn, a gap of at least 1224 h must be provided or the dye must be changed [22, 23]. Patency testing must be taken up in animals when other diagnostic procedures have yielded no conclusive information and the animal continues to evidence obscure infertility. Improvements in methods of assessment of fallopian tube functions in human medicine include hysterosalpingography and laparoscopic chromopertubation [261]. However, such techniques need the attention of veterinary practitioners.
Therapeutic Regimens The therapy of RB cows/buffaloes is deemed to be instituted only when oestrus detection and breeding (natural or AI) protocols are optimal. Therapies in a herd with suboptimal fertility constitute corrective measures to prevent/combat disease and/or deciency and reducing stress. Temporary replacement/change of the bull may take care of infertility because of the bull. The therapeutic regimens in a herd with reproductive failures must be aimed at the correction of the most probable causes. Treatment of individual cows/buffaloes at most situations remains difcult as a proportion of animals [22, 23] are always present with obscure reasons of poor fertility. Moreover, most diagnostic modalities described are largely unavailable to the treating clinician. With limited facilities the therapeutic approach usually must be aimed sequentially at (i) combating uterine infection (endometritis), (ii) correcting ovulatory disturbances, (iii) supplementing for luteal insufciency and (iv) improving management. When applied with sufcient caution, one or all of these approaches would culminate in the successful establishment of a pregnancy both in dairy cows and
10
buffaloes. The detailed therapeutic regimens used widely are described below.
Endometritis The clinical forms of bacterial complications of the bovine uterus have been described recently [123], and accordingly uterine infection that commonly occurs in repeat breeder cows and buffaloes falls under the denition of subclinical endometritis which clinically evidences after 8 weeks of parturition and a complete lack of cervical discharge with pathogenomic property [168, 169, 278]. Being a common cause of pregnancy failures, subclinical endometritis must always be suspected in repeat breeder cows and buffaloes, once we have assured the male factors (semen quality, bull). When infection with microbes in the uterus is suspected, there are a multitude of therapeutic approaches that have been used widely.
Antimicrobials Antiseptics such as dilute Lugols iodine [279] or povidone iodine [280, 281] have shown promise, to a limited extent, in therapy. Likewise, limited data show the success of the administration of antimicrobials in such cases ([282, 283]. Antimicrobial drugs administered the day after insemination to rid the uterus of organisms that might be detrimental to the survival of the conceptus have been commonly used in many countries [284]. Alternatively, antibiotics may be infused in the uterus for 35 days during oestrous and insemination is done in subsequent oestrus. Conicting reports depict the limited effect of treatment [285288] and a promising benet of treatment both in cattle [289295] and buffalo [296298]. The limitations of intrauterine therapy are development of drug resistance, inconsistent results and milk disposal after treatment that render such treatments uneconomical [299]. Moreover, the uterus seems to have a considerable capacity of spontaneous recovery, and a large proportion of animals probably do not require any therapy at all, especially under the aspect that some therapies are ineffective and might even cause more harm than benet [300]. A similar validation [301] suggests that when the endometritis is severe, intrauterine infusion offers benecial effects: however, in slight endometritis a similar treatment had a negative effect on reproductive efciency. Ott [543] considers that the result of intrauterine therapy is generally poor not only in repeat breeders, but also in animals moderately affected by endometritis. The in vitro sensitivity patterns of various antibiotics on the cervico-vaginal mucus collected from repeat breeder cows [302306] and buffaloes [307] have been described in an attempt to formulate the most effective antibiotic for therapy. However, until the specimens are collected directly from the uterus using specialized instrumentation
they do not represent the true picture of uterine infections as the mucus may be contaminated with microbes residing in the cervix and vagina and hence they cannot be recommended widely. When clinical or subclinical endometritis is suspected, the authors feel that, because of their low cost, properly administered antibiotics must be the clinicians rst choice if akes of pus are evident in the vagina or cervico-vaginal mucus as also suggested previously [308], and ushing of the uterus with normal saline as suggested previously [309, 310] must be considered when more of pus is evident clinically or when therapy alone with antibiotics fails. The route of administration for antibiotics in subclinical endometritis must be intrauterine as it leads to high concentrations of the drug in the uterine cavity and endometrium, and a relatively small amount is absorbed into the systemic circulation [311]. Systemic antibiotic administration should therefore be opted in treatment of more serious cases of metritis [312]. The most traditional antibiotic of choice has been oxytetracycline; however, because of its locally irritative character and increase in the minimal inhibitory concentrations (MIC) during the last decade, high doses (24 g/day, for 35 days) are required [123], which suggests opting for better alternatives. The expected in vivo efcacy of other traditional antibiotics (amoxicillin, and aminoglycosides) is questionable [282, 283, 313316]. The efcacy of nitrofurazones continues to be debatable, and only the clinical efcacy of uroquinolones has shown some advantage [317, 318] but the MIC of quinolones is not known. Likewise, the efcacy of penicillins given via an intrauterine route is doubtful [319]. Recently, the new (third and fourth) generation cephalosporins have shown efcacy against most uterine pathogens at low MIC values [316] and the rstgeneration cephalosporin (cephapirin) is recommended for intrauterine use [166, 197, 320] as the drug of choice for subclinical endometritis [123]. It is suggested under most bovine and bubaline situations to combine an antibiotic with an imidazole derivative (metronidazole or tinidazole) to take care of anaerobic microbes and protozoa that may inadvertently be present [318]. Addition of antifungal agents as suggested previously [321323] must be considered when the endometritis turns out to be chronic after too much of therapy with antimicrobials. Plenty of alternative therapies for treatment of endometritis have appeared in the recent past. Of these, the most potent and safe approach appears to be the use of prostglandins (PG) [288, 324, 325]. The uterus has an increased inux of PMNs, an increased blood supply, increased mucus production [326] and enhanced uterine production of leukotriene B4 during the oestrus period because of increase in proinamatory cytokines stimulated by PG. The immune functions of the uterus are thus enhanced [327]. Hence, returning cows and buffaloes to oestrus at short intervals would lead to endogenous clearance of microbes and cure from endometritis. This
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can usually be achieved by injecting a PG from 510 days of oestrus alone [5, 288, 301] or preceded by a uterine lavage [310]. In clinical practice it is sometimes recommended to keep the cow/buffalo indoors at 23 oestruses to avoid matings and this works in recovery from endometritis well, but suffers from the longer time interval that passes by before the animal conceives and it is always better to use prostaglandins.
Immunomodulators In the recent past, several therapies alternative to the use of antibiotics have been suggested for the therapy of endometritis. The intrauterine infusion of immunomodulators such as E. coli lipopolysaccarides (endotoxin) [195, 328335], oyster glycogen [209, 283, 336338], infusion of serum, plasma or hyperimmune serum [283, 331, 334] or leukotriene B4 [339] has been reported widely. These immunomodulators act as a chemoattractant to the PMNs through stimulation of interleukins [340] produced by monocytes and macrophages. The PMNs, blood monocytes and macrophages are regarded as the professional phagocytes in the cellular defences against pathogenic micro-organisms [330]. After experimental intrauterine infection, the PMN population within the uterine lumen usually increases [341, 342]. A single intrauterine infusion of 100 mg of E. coli LPS dissolved in 20 ml of phosphate-buffered saline (PBS) results in increase in the uterine neutrophils (of up to 80%) within 6 h, which remains for 72 h [232, 233]. Likewise, 0.110% oyster glycogen (OG) (usually 500 mg) dissolved in 60 ml of vehicle or 30 nmol/l of leukotriene B4 increases the PMN concentration within 1224 h of administration [195, 339]. Within 72 h of administration of either LPS or OG, the denuded epithelium of endometritis-affected crossbred cows was reduced and the psuedostratication of uterine endometrium was completely cured [337]. However, LPS was found to cure all types of endometritis except the chronic type with cystic dilatation of endometrial glands [337]. Using these treatments the endometritis would usually be cured and cows can be inseminated at subsequent oestrus. However, LPS is known to suppress follicular growth, decrease oestradiol production and delay the LH surge and ovulation [343, 344], and thus the subsequent cycle may be delayed. Addition of a small amount of autologous serum or plasma (50100 ml for 23 days) to uterine secretions increases the opsonizing capacity and signicantly enhances the phagocytic ability of PMNs [283]. Besides the use of immunomodulators, some other therapies suggested for resolving endometritis include the use of enzymes and antioxidants. Enzymes like trypsin, chymotrypsin and papain when infused into the uterus resulted in a cure rate of 59.7% (Revealed by absence of vaginal discharge at re-examination); however, the conception rates were suboptimal [345]. Another enzyme
that has been tried is lysozyme [346] with a good success rate. Some medicaments like 4 mM taurine and 50 mM fructose in PBS [347] and ascorbic acid (vitamin C) have been used for intrauterine infusion in order to change the uterine pH prior to insemination, and act as an antioxidant (G.N. Purohit, unpublished work). Antioxidants such as vitamins C and E are known to modulate the oxidative stress and reduce the endometrial damage both at the biochemical and histological levels [348]. The stressors, free radicals and b-endorphins were higher in repeat breeder cows in a recent study [349]. However, whether the benet of intrauterine infusion of antioxidants like vitamin C in repeat breeder cows in our clinical trials was by resolving endometrial damage or by reducing the concentrations of b-endorphins and free radicals generated because of stress remains to be validated. Moreover, frozen semen might have damaged sperms or sperms with altered function because of the reactive oxygen species generated during the freeze thaw process [350] as semen has little antioxidants to protect them [351]. An infusion of antioxidants before AI might reduce the uterine luminal reactive oxygen species and the b-endorphin that might reduce the functional competence of frozen spermatozoa, which might not have been offered by parentral administration of antioxidants. A positive correlation has been previously observed between lipid peroxidation levels of plasma and cervical mucus of cows [352].
Correction of Ovarian Dysfunction Ovulatory disturbances culminating in RB cows and buffaloes include anovulation and ovarian cysts. The effects of both delayed ovulation and anovulation in terms of end result appear similar: pregnancy failure and hence RB. The patterns of follicular development during periods of anovulation have been described for cattle [353]. Delayed ovulation results in poor fertility [102]. The underlying physiology of anovulation seems to be a lack of a preovulatory surge in response to the high, oestrual concentrations of oestradiol [105], presumably because of lack of progesterone priming of hypothalamus or a multitude of other factors. Anovulation conditions with growth of follicles to deviation but not to ovulatory size may be because of undernutrition, suckling or diseased condition. Another reason for anovulation may be the presence of suprabasal progesterone concentrations during oestrus, which has an inhibitory effect on the positive feedback of high oestradiol concentrations on the hypothalamus, resulting in high LH pulse frequency and effects on follicular growth [354]. Spontaneously occurring delayed ovulation or anovulatory conditions have been reported in connection with RB in postpartum cows [103, 104]. Several studies have reported development of persistently dominant follicles subsequent to pharmacologically induced suprabasal progesterone concentrations in dairy cows, with inhibitory effects on oestrus signs and
12
LH release in a dose-dependent manner [355358]. In cases of anovulation, there is some evidence for an absent or decient LH surge trigger mechanism: therefore, GnRH treatment can stimulate ovulation and, in some cases, result in increased pregnancy rates [359, 360]. It thus appears that many anovulating events are primarily the result not of ovarian disorders but rather of decient hypothalamic function. Different types of anovulatory conditions have been described in cattle [105]: however, clinically, therapies to stimulate ovulation in dairy cows and buffaloes with delayed ovulation/anovulation essentially remain the same and include administration of either hCG (15005000 IU, intravenous, or 500010 000 IU, intramuscularly) [36, 114, 361364] or 100 mg of GnRH [359, 360, 365377] or hMG [378]. These therapies usually evoke LH release [360, 379] but the CL formed by hCG injection had a shorter life span [380, 381] and 67% of the induced short cycles were followed by a return to acyclicity [382, 383]. Alternatives to these well-known therapies include administration of glucose and insulin, prostaglandins, metformin, antiprolactins and clomifene. The LH surge is known to be complex and affected by an interplay of various endocrine, neurocrine, metabolic and cellular events. Low levels of glucose, insulin and insulin-like growth factors all affect the LH surge [59]. It has been reported that intramuscular (IM) administration of 0.2 IU/kg bovine insulin to dairy cows on days 8, 9 and 10 of oestrus resulted in increased concentrations of progesterone in treated cows and high levels of insulin and glucose in cows that conceived than those that did not conceive [384]. The authors have attempted treatment of dairy cows with delayed ovulation by administration of 500 ml of 25% dextrose at oestrus along with insulin (5 ml of bovine insulin). In preliminary trials, 30 of the 50 cows treated with such a treatment had timely ovulation and conceived subsequently. However, such therapies need to be validated further in planned studies before they can be recommended. Prostaglandins are known to be involved in the ovulation process as they increase the intrafollicular uid pressure and follicle wall thinning [327] and as such can be used as agents to facilitate ovulation. Moreover, the presence of luteal tissue at the time of nal follicle growth pez-Gatius et al. [386] may hamper ovulation [385]. Lo noticed that intravenous cloprostenol at AI promoted ovulation in repeat breeder cows and cows with stress. Treatment of ovarian cysts with GnRH is also known to yield a better therapeutic outcome when combined with a simultaneous prostaglandin [387] administration. Clomiphene citrate, an antioestrogen, is known to exert direct antiovulatory and oestrogen antagonistic actions in rats [388]. Tamoxifen and clomifen are mixed antagonistagonists of oestrogen action and belong to the group of type I antioestrogens [389]. Type I oestrogen antagonists partially inhibit the action of agonists, but due to their own inherent weak agonistic properties, they also induce, to some extent, oestrogenic responses. The
degree of agonistic or antagonistic activity depends on the species, organ, tissue or cell type that is being examined [390]. In women, clomiphene is a well-known pharmaceutical agent for ovulation induction in patients with polycystic ovarian disease [391393]. It may exert action directly on pituitary gland to augment oestrogen-induced LH release [394]. Only sparse reports are available for the use of this drug in animal studies. A dose of 300 mg of clomiphene citrate administered to cows after a 1% copper sulphate drench has been suggested for the treatment of anoestrus in cows and buffaloes [395398] and for the treatment of RB [399] and cystic ovarian disease [400] in cows. In known cases of ovulatory disturbances in cattle and buffaloes, clomiphene should thus be started preferably 1 day before oestrous (300 mg orally after copper sulphate at 12 h intervals) until the onset of oestrous. The up-regulation of receptors that follows a down-regulation would facilitate LH release and ovulation. It has been suggested in human therapy that in clomiphene-resistant women, metformin, an insulin sensitizer, combined with clomiphene, could be a better option for ovulation induction in patients with polycystic ovarian syndrome [391, 393, 401] as insulin sensitizers improve hyperinsulinaemia and hyperandrogenism in treated women [402]. However, such therapeutic agents need validation in the ovulation induction programmes for cattle, and buffaloes as well. The authors have clinically used 20004000 mg of metformin given orally to RB cows, but the outcome is unknown, at the moment. Yet another medicament, the antiprolactin bromocryptine have been suggested by some clinicians to help in ovulation induction in RB cows (P.K. Pareek, personal communication) with suggestions of 10 mg given orally 12 h before and at the time of AI. In trials on ewes, 0.61.2 mg of bromocryptine administered orally for 312 days decreased prolactin but did not affect FSH, the mean time of LH preovulatory surge or LH concentration in LH surge [403, 404]. The administration of antiprolactins in ovulation induction therefore remains questionable. Besides ovulatory failures resulting in failure of conception because of ovulationinsemination asynchrony, the most common ovulatory disturbance recognized in dairy cattle is the cystic ovarian disease. The problem is less known (with an incidence of 0.24%) clinically in the buffalo although reported in many studies [405409]. The description of the presence of follicular cysts is presumptive as signs of nymphomania, mucometra, and frequent oestrous have never been recorded in buffaloes with cystic ovaries [16]. Cows with single or multiple follicular ovarian cysts and normal oestrus cycle lengths are often presented to the clinician with a history of RB. When follicular cysts persist for prolonged periods in dairy cows, endometrial gland hypertrophy and pathologies in the uterus many a times culminate in clinical mucometra with normal oestrus cycle lengths. The therapeutic management of ovarian cysts in dairy cattle has been reviewed previously [108, 109, 410,
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411]. It has been suggested previously that treatments are only temporary solutions and it is better to select cows against cysts [108]. The therapies suggested include administration of a single IM 200 mg injection of progesterone in oil [412] or insertion of intravaginal progestagen implants [413, 414], 100 mg of GnRH [106, 107, 369, 415417], hCG [20, 106, 418420] or prostaglandins [106, 410, 411], depending on the type of cyst. More recent literature suggests a combination of these therapeutic agents [387, 421423]. Accordingly, GnRH or hCG treatments are followed by prostaglandin treatments after 810 days. The rationale for this combination appears to be luteolysis of leutinized tissue formed by administration of either GnRH, hCG or progesterone. The complex physiology involved in the formation of cysts active at the hypothalamus, pituitary, ovary and metabolic levels and the follicle turnover mechanisms, however, render the treatments suboptimal with a tendency of cysts to reform some time after disappearance. The Ovsynch protocol suggested and used widely [424, 425] utilizes the administration of GnRH (100 mg) on day 0, followed by prostaglandin on day 7, GnRH (100 mg) on day 9 and AI 1620 h later. The second dose of GnRH assures pez-Gatius and ovulation of the newly formed follicle. Lo pez-Be jar [387] were of the opinion that a dose of Lo 500 mg of prostaglandin administered along with GnRH on the rst day of treatment followed by a second dose of prostaglandin 14 days later resulted in a lower cyst persistence and higher ovulation rate compared with when GnRH was given alone on the rst day of treatment. Aspiration of follicular uid from follicular cysts using transvaginal ultrasound-guided aspiration has been reported to be a new concept in treatment of follicular ovarian cysts in dairy cows [426]. Whatever the therapy adopted for treatment of ovarian cysts in dairy cows, there is a tendency of cysts to reform, and the difculty in such a clinical condition lies not in the resolving of the cyst but in the attainment of a successful pregnancy, which is extremely difcult when cows develop clinical mucometra. Regimens suggested to resolve mucometra include oral (310 g of potassium iodide for 510 days) [106, 419, 427, 428] or injectable administration of elemental iodine [281] or uterine lavage [309]. However, such therapies are often unpredictable. Much has been written regarding suprabasal progesterone at oestrus in dairy cows and low conception [92, 95, 96, 235, 236, 429, 430] or ovarian cyst formation [431434]. However, little has been suggested on correction of such a high progesterone concentration at oestrus. The extra progesterone thought to be of adrenal origin [435, 436] could not be conrmed to be originating from the adrenals in studies by Bage et al. [98]. However, environmental or social stresses were postulated to originate adrenal stimulation with resultant deviant hormones and RB. The diet and milk production both can also alter the progesterone [437] and, as such, the only probable preventive measure to reduce suprabasal
progesterone would be monitoring the diet and reducing stress. Although deciency in positive feedback of oestrogen to the hypothalamus, leading to a lack of LH surge with a resultant anovulation, is the widely accepted cause for ovarian cysts, a more recent postulation for ovarian cyst formation is the delay (or absence) of the degeneration system of the unovulated follicle [438]. Some new concepts in the formation of ovarian cysts in dairy cattle include a low insulin concentration [432], an increase in FSH following a reduction in inhibin secretion [439], a decrease in IGF-1 in follicular uid [440], altered oestrogen receptors [441], and alterations in expression of cytoskeletal proteins in follicles [442]; however, such insights into the formation of cysts do not necessarily affect therapeutic regimens.
Luteal Insufciency Luteal inadequacy due to diminished response to the circulating luteotrophic hormones [443] leads to insufcient progesterone production during the luteal phase after breeding, and could be the cause of embryonic death [444]. The serum progesterone is known to be altered in RB cows [96, 238, 240, 445448] and buffaloes [78, 239, 443]. Shelton et al. [443] argue that luteal inadequacy, caused by a diminished response to circulating luteotrophic hormones, may contribute to embryo mortality in subfertile cows. Early in the luteal phase the progesterone down-regulates the oxytocin receptors (OTRs) for at least 10 days, thus preventing premature luteolysis [29]. The secretion of antiluteolysin factor IFN-t and bovine trophoblastic protein-1 (bTP-1) around day 1516 post ovulation mainly depends on progesterone concentration around day 45 post ovulation [449]. Moreover, a conceptus has to be around 15 mm long to secrete IFN-t, and its growth is largely dependent on progesterone levels [450]. A low progesterone level has been shown to be signicantly related to reduced production of IFN-t by bovine embryos recovered on day 16 of pregnancy [451]. The most critical period for embryo survival may be around day 56 post insemination when the embryo descends from the oviduct and enters the uterine lumen. During this period, progesterone concentrations start rising and thus any delay in the rise and/or a low luteal phase progesterone concentration can cause poor embryo development and hence embryonic death due to a suboptimal uterine environment on account of low progesterone [452]. Inadequacies of luteal tissue formation could also arise on account of poor development of a preovulatory follicle [453], presumably because of low IGF-1 [454]. Nutritional inadequacies can result in deciencies of growth hormone and/or insulin with resultant low IGF-1 secretion [455]. Likewise, low progesterone in buffaloes is also a result of breeding season, which causes inadequate functioning of CL [377, 456], possibly because of high prolactin [7].
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The therapy for luteal insufciency is based either on evaluation of blood and milk progesterone on day 5 post insemination, or solely based on presumptions. Therapeutic regimens include administration of one of the following: (i) GnRH at the time of insemination [457] or day 1113 post AI [373]. The mechanics of GnRH action on luteal insufciency prevention are explained elsewhere [29, 458]. (ii) hCG on day 47 [459] or day 1516 post AI [460] as this period coincides with the presence of dominant follicles in cows having three follicular waves. Peters [29] suggested that administration of hCG on day 11 and 13 is most benecial as around this period maternal recognition of pregnancy occurs. The benet of hCG or GnRH therapy was postulated to be because of formation of an accessory corpora lutea [29, 458]. However, a more recent study showed no effect of double ovulation (in 35% of animals) on luteal function or plasma progesterone concentrations [232]. (iii) Progesterone supplemented as a single IM injection (500 mg) on day 5 post AI [461], chlormadinone daily oral feeding (10 mg) from day 1423 [462] or progestagen vaginal implants from day 512 of AI [362, 452, 463465]. Progesterone supplementation advances the luteolytic signal and increases embryonic growth [466] and thus increases pregnancy rates when given during the rst week post AI, the most critical period of embryomaternal interactions [4]. (iv) Recombinant bovine somatotropin (500 mg SC) at the time of oestrus and 10 days later signicantly increases the conception rate because of increase in circulating progesterone [467]. Besides the above therapies, some other less common therapies suggested include feeding of linoleic acid, as it is an inhibitor of prostaglandin synthetase enzyme and thus delays premature luteolysis and enhances luteal function, and feeding of sh oil, as it contains docosahexanoic acid and eicosapentaenoic acid, both of which have antiluteolytic properties [468].
Management Strategies The overall management of dairy cows and buffaloes is important as it affects the fertility. Of consideration are nutrition, timing of insemination and periparturient disease.
management of the dairy cow, particularly before and after calving, has been considered a key driver of infertility [81]. Some of the signicant reviews appearing recently include reviews on the effects of macro- and microminerals during the periparturient period [82], the impact of controlled nutrition during the dry period [83], the effect of rumen degradable proteins [84], and embryo survival in dairy cows managed under pastoral conditions [33]. It appears from all these and other published data that for todays high-producing dairy cows, fertility is in general heading towards a decline [6] although, for the parous cow, feeding during the dry period and postpartum appears to be crucial in maintaining high fertility. Inadequate body condition postpartum has a greater impact on the probability of conception and embryonic losses [496]. While poor nutrition during the dry and early postpartum period results in reduced glucose, insulin, insulin-like growth factor (IGF-1) and low LH pulse frequency with concomitant increases in b-hydroxybutyrate, non-esteried fatty acids and negative energy balance all having negative effects on the probability of conception. Conversely, high nutrition can also increase the metabolic clearance rate of steroid hormones such as progesterone and oestradiol, and high rumen degradable proteins can raise the blood urea nitrogen. All these can impair conception and embryo survival. However, the impacts of nutrition on fertility appear to be complex, and recommendations for formulating effective dietary strategies to improve conception rates and prevent embryonic losses during the more crucial stages therefore appear to be difcult. In general however, it has been recommended that cows must not lose excessive body condition postpartum, and should not be fed more than 10% of rumen degradable protein. A balanced feed during the dry period must therefore comprise a low-energy high-bre ration containing high levels of chopped straw. These recommendations, however, do not point out the possible deciencies in clinical cases of RB cows and buffaloes, which may have one or multiple deciencies or excesses. It is the authors presumptive view that clinical cases suffering from the RB syndrome at many locations suffer from multiple deciencies, especially those of glucose, vitamins such as A, E, and C and minerals like phosphorus, calcium and selenium and, as such, animals must be supplemented with these nutrients by oral or injectable supplementation. Some of the published literature does not concur with the authors view [497, 498], while other reports do [494, 499504], essentially because most of these trials were performed on wellmanaged herds and not on individual cows or buffaloes.
Improving the Timing and Technique of Insemination Improving Nutritional Imbalances The effects of nutrition on fertility in dairy cattle have been extensively reviewed recently. Poor nutritional Much improvement can be expected by improving the timing of insemination essentially by appropriate oestrus detection. A sizeable proportion of cows evidence
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prolonged oestrus periods and such animals pose problems to time insemination [505]. Multiple inseminations or ovulation induction treatments are suggested in such animals. Heifers frequently evidence short oestrus periods and it is suggested to time insemination slightly earlier. Buffaloes pose a greater difculty in oestrus detection and sub-oestrus is frequent, hence timing inseminations becomes difcult. Vaginal electrical resistance measurements have been suggested for oestrus detection and timing insemination both in cattle [506] and buffalo [507, 508] with limited success. Likewise, the use of pedometers [544545] and radio-telemetric devices [546547] has been suggested to improve oestrus detection and, hence, timing of insemination. The usual timeframe suggested for timing inseminations in cattle [509] have been suggested to be repeated twice at 12 h interval in the buffalo for optimum conception rates [11]. It has previously been suggested for cows that if onset of oestrus is unknown (which is frequent for animals submitted to inseminators under most situations), inseminations should be performed within 6 h of initial observation of oestrus [548] because 24.1% of cows have oestrus of low intensity and short duration [547]. Fixed-time AI subsequent to an oestrus synchronization protocol has been shown to improve fertility of dairy [549] and beef [550551] cattle. AI is usually scheduled 6072 h of a PG injection, and such protocols signicantly improved fertility of cows suffering from heat stress [549], as heat stress is known to decrease the intensity and duration of oestrus expression and increase the incidence of anoestrus and silent ovulation [552]. However, the use of xed time AI in RB animals appears to offer little advantage, and other important aspects mentioned elsewhere in this review with often overlapping effects are of greater signicance. The usual site of insemination suggested both for cattle and buffaloes is the mid-cervix; however, some reports depict benets of insemination in the uterine horns [510 513], probably because the functional sperm reservoir near ovulation is the uterine portion of the oviducts and not in the cervix [514]. A rigid insemination device, the Ghent device was reported to be developed for deep intrauterine insemination at the uterotubal junction in dairy cattle [515]. However using the usual AI gun or the Ghent device or reducing the sperm numbers from 12 to 4 millions had no effect on pregnancy rates with either of the methods [511, 515]. In trials during summer months no benet of depositing semen in the middle of the uterine horn or uterotubal junction using low or standard dose of sperms was observed on the pregnancy rates in cattle [512, 516]. Likewise, Momont et al. [517] concluded from their trials in cattle that placement of semen in one horn of the uterus does not appear to be a cause of decreased or increased pregnancy rates with AI. In buffaloes, Zicarelli et al. [518] had contrarily suggested that inadvertent deposition of semen in the uterine horns of buffalo due to the small size of the uterine body could be the reason for low conception rates. Moreover, according
to Vale [49] a pregnancy rate higher than 50% can be considered good after insemination with frozen thawed buffalo bull semen. In the authors view, deposition of semen in the body of the uterus offers a distinct advantage in improving the conception rates to AI both in cattle and buffaloes, compared with when it is deposited in the midcervix. Errors in the preparation of the AI gun or those in the upkeep of frozen/liquid semen can contribute to conception failures and hence must be viewed seriously. A recent report depicts that the conception rates to articial insemination improved by 527% over many Asian countries when the personnel involved in the AI were given various levels of training [12].
Avoiding Periparturient Disease The role of prevention of problems in the periparturient period, in particular hypocalcaemia, mastitis and retained placenta, has been stressed in a recent review [82] as all are known to have a negative impact on the subsequent fertility of the cow. Likewise, metabolic diseases during the postpartum period, such as ketosis and acidosis, or parturient problems, such as dystocia, predispose cows/ buffaloes to development of postpartum uterine diseases such as endometritis with the result of more services per conception [81]. The approaches suggested to reduce the incidence of these disorders to some extent include the feeding of anionic salts in combination with adequate calcium and magnesium [82] during the dry period and feeding of high-bre low-energy chopped straw during the dry period [83]. However, although parturient problems appear to be unavoidable, stress must be attached to parturient hygiene. Many locations where cattle and buffaloes are raised suffer from extremely poor hygiene. Moreover, often animals are referred for therapy only when they have a reduced feed intake/milk production or develop serious clinical signs. Coupled with this is the fact that farmers attending calving or manually removing plancentas often handle animals without any sanitary measures. These practices are likely to precipitate low conception levels postpartum presumably because of lowgrade infection or damaged genitalia. It is therefore important to educate farmers regarding the possible consequences of the poor hygiene at calving and post partum.
Reducing Stress Stress appears to play an important role in the modulation of various biological events including reproduction. The role of various types of stress because of disease, inadequate nutrition, high production, social factors and environment on reproduction has been explained previously [154]. It is nearly impossible to avoid all forms of stress in dairy cows and buffaloes, but when animals
16
Repeat breeder cow/buffalo
Exclude effects of season
Herd
Individual 1. Investigate for abnormalities of genital organs like ovaro-bursal adhesions, cystic ovaries, tumours, stenosis, etc. 2. Investigate for subclinical endometritis. When no tests possible, treat on presumptions if there is a history of periparturient disease. 3. Monitor ovulations/oestrus cycle length (i) Provide ovulation induction treatments at AI (ii) Repeat AI/consider I/U AI 4. If animals do not settle, treat for luteal insufficiency. 5. Supplement with vitamins A,E and C and Ca, P and Se. 6. PSP dye test if both fallopian tubes occluded. Exclude such animals. 7. Cytogenetic-karyotyping
Female Investigate and advise 1. Nutrition (preparturient) 2. Collect samples for investigation of infectious disease 3. Reduce stress 4. Metabolic profiles
Male
AI 1. Evaluate semen and AI techniques
Natural mating 1. Infectious disease (i) Trichomonas (ii) Campylobacter 2. Semen evaluation 3. Age of bull
Figure 1 Diagnostic approaches for repeat breeding
require more number of services per conception, attempts must be oriented to minimize stress associated with environment. Cooling of cows/buffaloes during hot summer months by showering of water is known to improve fertility. Likewise provision of sufcient shade and wallowing space essentially improves conception rates in buffaloes which inherently have a poor thermoregulatory mechanism.
Imunoinfertility It has since long been postulated that sperm, when deposited in the female tract, can act as an antigen and evoke production of antibodies leading to immunoinfertility [242]. Antibodies to sperm may appear both in the blood and in genital tract secretions of human females [242]. Both antisperm IgG and IgA have been isolated from bovine [146] and human subjects [245, 246]. These antisperm antibodies usually reduce the sperm penetration of cervical mucus with immobilizing activity of sperms in cervical mucus of woman [244246]. Although agglutination of bovine sperm with cervical mucus has been reported through in vitro studies [129, 130, 146], in vivo studies have not conrmed their signicance [131, 138]; therefore it appears that immunoinfertility is more a human concern, and in cows and buffaloes the presence of such a phenomenon continues to be anecdotal. Studies by Tripathi et al. [147] found that of the 17 sperm-specic polypeptides detected to be reactive with antisperm antibodies, only two were reactive with sera from presumptive immunoinfertile cows. The different methods of cervical mucus penetration assays used for bovine
examinations are more suitable to evaluate motility of sperm than to predict potential fertility [139, 248, 249] and hence their use to predict infertility of immunogenic origin similar to that used in human subjects is suboptimal. Therapies of immunogenic infertility in animal subjects appear to be simple; changing the semen or intrauterine insemination would help taking care of any such problem. Therapies in human subjects are beyond the scope of the present review and mostly difcult as sperms to which antibodies are produced continue to travel through the female genital tract. A few of the approaches include administration of vitamins C and E and dexamethasone [247, 519] and intrauterine insemination of vitamin C (G.N. Purohit, unpublished work) with little success.
Miscellaneous Therapies Despite all efforts of therapy, a proportion of RB cows and buffaloes would continue to be infertile for prolonged periods and they are described to have infertility of unknown origin [22, 23]; such an infertility should better be designated as idiopathic. Therapy of such idiopathic infertility is seldom possible. Some of the less common therapies described for cows suffering from the RB syndrome include acupuncture therapy [520], intraperitoneal insemination [521], use of herbal drugs [522, 523] and embryo transfer at 78 days of oestrus with or without AI at oestrus [524]. Such therapies, however, have little to offer in improvement of the condition. Cows that gain excess of body fat are a classic example of idiopathic infertility. Such cows, when made to lose weight by severe diet restriction, often conceive. It is difcult to
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comment on such infrequent therapeutic approaches as their results are inconsistent. Treatment Approaches The approach to therapeutics in herds or individual animals is signicantly different. When low fertility to either AI or natural services is a problem in herds, investigations must be made on the presence of infectious diseases such as Campylobacteriosis, by random collection of specimens. Moreover, when natural service is being used, the bull used must be investigated. Specimens of blood should be collected to evaluate metabolic proles. Approaches of feeding suggested during the dry period and the post parturient period can help to correct some nutritional inadequacies and those on combating uterine infection can reduce fertilization failures or embryonic mortalities. When faced with therapy of individual animals, the approach should be rst to treat uterine infections and then treat ovulatory disturbances or corpus luteum inadequacies (Figure 1). In spite of all these therapies, a small proportion of cows/buffaloes would have infertility of unknown origin and it is still extremely difcult to delineate or treat such obscure infertility. Conclusion It can be concluded that diagnosis and therapy of RB continues to be difcult, but when investigating and treating individual cows/buffaloes, a systematic approach of combating uterine infection and correcting ovarian dysfunction or luteal insufciency would result in a majority of animals to conceive provided the management and breeding techniques are optimal. The feeding of highproducing cows, especially during the periparturient period, minimizing stress and parturition hygiene are crucial to obtain high-fertility postpartum, and dairy farmers must be educated in this regard. While making selection of cows for high production, stress must now also be laid on high fertility. Diagnostic approaches such as hysteroscopy must be stimulated to widen our knowledge of the inner surfaces of the genital tract.
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Acknowledgements I am grateful to Dr Vijay Kushwaha, Dr Manish Garg and Dr Arvind Sharma for the endless help they extended in the preparation of this paper. References
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Repeat Breeding in Cattle and Buffaloes

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CAB Reviews: Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources 2008 3, No.

CAB International 2008 (Online ISSN 1749-8848)

Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources

Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources

Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources

time-consuming and often inconclusive in diagnosing the cause of RB in an individual animal.

Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources

Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources

Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources

Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources

Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources

Repeat breeder cow/buffalo

Exclude effects of season

AI 1. Evaluate semen and AI techniques

Figure 1 Diagnostic approaches for repeat breeding

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