The natural resistance of cattle to artificial infection with Cowdria ruminantium: the role played by conglutinin

The conglutinin titres of year-old Bonsmara-cross cattle infected with Cowdria ruminantium were inversely proportional to the severity of the reactions elicited by the infection. There was no correlation, however, between conglutinin levels of 8-month-old calves of the same breed, sex and origin and their susceptibility to heartwater. The role possibly played by conglutinin in the non-specific resistance of cattle to heartwater and in the epidemiology of the disease is discussed.     

The immunization of calves against heartwater: subsequent immunity both in the absence and presence of natural tick challenge

Cattle, vaccinated as calves with Cowdria ruminantium-infected tick stabilate, were challenged 6, 12 and 24 months later. In the absence of tick challenge, vaccination of calves induced a partial immunity against subsequent challenge at 12 and 24 months. In animals exposed to ticks, the resistance was no better than that of control, unvaccinated cattle. When they were challenged at 6 months of age there was no difference between vaccinated and unvaccinated calves, either in the absence or presence of tick challenge, and all the animals manifested a high degree of natural resistance. This study therefore suggests that the value of vaccinating Afrikander-cross calves in heartwater endemic areas should be further investigated. The indirect fluorescent antibody (IFA) test proved to be a valuable means of monitoring the serological response of vaccinated animals and detecting the sero-conversion of animals exposed to tick infection. On one hand, there was good correlation between the febrile reaction and the results of the IFA test on the sera of vaccinated and control cattle challenged with the heartwater agent, in that all sero-positive animals were resistant to challenge. On the other hand, though, a considerable percentage of the animals that were serologically negative were also resistant to challenge.     

Mortality percentages related to heartwater and the economic impact of heartwater disease on large-scale commercial farms in Zimbabwe

A postal questionnaire asking for data concerning heartwater deaths was sent to all large-scale commercial farmers in Zimbabwe's lowveld, where heartwater is endemic. One hundred and fifty returns (24% return rate) provided usable data, with 85 farms (57% of sample) reporting heartwater deaths in 1990/91. The median heartwater mortality percentage for calves (0–12 months) was 2.3%. This was statistically significantly different (Wilcoxon signed rank test, P < 0.005) from the median mortality percentage of 0.6% for cattle 13 months and older. Heartwater accounted for 51% of all mortalities on farms reporting heartwater deaths. There were no meaningful correlations between farm or herd size and heartwater mortality percentages. A plot of the farms reporting heartwater deaths found that the heartwater endemic area in 1991 is essentially unchanged from 1979. Inspection of the geographic distribution of farms reporting heartwater showed no obvious patterns, and distances from communal land farms were not significantly correlated to heartwater mortality percentages. An economic model found that the annual financial impact of heartwater on large-scale commercial farms in Zimbabwe to be some Z$19.4 million (1991 prices). The upper bound, that allowed for inflation since 1991, was Z$33.4 million. The lower bound estimate was Z$8.1 million (Z$3 = US$1, 1991 rates). Eighty-nine percent of these costs can be attributed to the cost of dipping. Since it has been shown experimentally that the carrier state of heartwater may not affect the productivity of cattle, intensive dipping may not be the best means of controlling heartwater.     

The non-specific resistance of cattle to heartwater

A distinction is drawn between the non-specific resistance to heartwater of very young calves and that of cattle older than a year. The suckling-calf resistance is at its greatest during the first few weeks after birth and then gradually declines during the next 6-8 months. Aspects of its possible mechanism and significance are discussed. The role played by conglutinin in the natural resistance of older cattle to heartwater and in the epidemiology of the disease in discussed.     

The treatment of heartwater

This paper reviews the available literature on the treatment of heartwater and draws comparisons with results obtained from recent work on experimentally-induced heartwater in sheep, calves and mice.     

Heartwater. An overview of the clinical signs, susceptibility and differential diagnoses of the disease in domestic ruminants

Heartwater is a frequently fatal tick-borne disease of ruminants caused by Cowdria ruminantium. In domestic ruminants the incubation period varies considerably and depends on the route of infection, virulence of the isolate and amount of infective material administered. Adult cattle of all breeds appear to be equally susceptible to heartwater. It is generally accepted that calves up to the age of 3 weeks have a high degree of natural resistance which is not related to the immune status of the dam. Nervous symptoms are frequently seen in animals affected by the peracute and acute forms of heartwater and can easily be confused with similar signs caused by infectious conditions, toxic plants, acaricide and heavy metal poisonings.     

An in vivo comparison of the efficacy of the heartwater blood and ground-up tick suspension vaccines in calves

Two groups of calves were respectively immunized with heartwater blood (BV) and ground-up tick suspension (GUTS) vaccine. A third group was left unimmunized as controls. No difference in the immune status conferred could be demonstrated between the 2 vaccines at 6 months and 12 months challenge after vaccination. An index, based on the rectal temperature before and during the reaction, was calculated as an aid in evaluating the data. In practice, the evaluation of heartwater vaccination by challenge is more effective at 12 months than at 6 months after vaccination. No effective difference was demonstrated between the 2 vaccines in their immunizing efficacy.     

Use of delayed-type hypersensitivity test to diagnose brucellosis in calves born to infected dams

Calves (n = 2) born to dams with experimentally induced brucellosis, and calves (n = 4) born to dams with naturally occurring infection were examined by the delayed-type hypersensitivity (DTH) test for possible B. abortus infection. The results were compared with the serum agglutination test, complement fixation test, and Coombs test. Five calves were nursed by their dams for 8-10 weeks after birth. One calf was separated from its dam and fed artificial milk. Three to five months after birth, four calves tested seropositive in the serologic tests. Antibodies were detected in one calf as early as 1 week after birth. The calf fed on artificial milk was seronegative 4-5 weeks after birth. All calves reacted to the DTH test antigen from week 12 until the end of the experiment, even though serologic tests were negative. We conclude that the DTH test is a valuable technique for diagnosing Brucella in calves born to infected dams.     

T cell-mediated immunity to Cowdria ruminantium in mice: the protective role of Lyt-2+ T cells

The inability of athymic nude mice to make a drug-aided recovery from infection with either the Kümm or the Welgevonden stocks of Cowdria ruminantium and their inability to mount an immune response, suggest that immunity in heartwater is cell-mediated. The adoptive transfer of immunity with the spleen cells of mice immune to the Welgevonden stock is supportive evidence. Immune spleen cells depleted of Lyt-2+ T cells are unable to confer resistance to challenge to recipient mice, whereas the depletion of L3T4+ T cells had no effect on the protection conferred by immune spleen cells. This is conclusive evidence that immunity in heartwater is largely cell-mediated. Immune serum, C. ruminantium and complement incubated in the presence of mouse peritoneal macrophages, inhibits the infectivity of the heartwater agent, but not in the absence of macrophages. The decreased resistance to challenge of immune mice treated with gloxazone adds further support to the concept that in heartwater persistence of C. ruminantium in the host is associated with immunity.     

Heartwater in Nigeria

A study was carried out to determine which materials from animals dying or dead of heartwater could initiate the disease in susceptible goats, using the intravenous and subcutaneous routes. C. ruminantium was consistently isolated by intravenous injection of the whole blood or of lung macrophages and by subcutaneous injection of brain homogenate. In animals dead of heartwater, it appeared that isolation of the organism was achieved only when extensive post-mortem autolysis had not supervened. Experiments with blood fractions showed that leucocytic and plasma fractions of infective blood transmitted heartwater; the erythrocytic fraction consistently failed to induce an infection.     

Problems with the interpretation of epidemiological data in heartwater: a study on 23 farms

In an epidemiological study undertaken on 23 farms where heartwater occurs endemically, it was found that on an overall average, antibodies to Cowdria ruminantium were detected in 64.3% of the cattle, 6 adult Amblyomma hebraeum ticks were counted per animal and 7.0% of ticks were infected with the heartwater agent. It was found that the seropositivity of the animals was determined largely by the tick loads to which they were subjected and that the influence of the tick C. ruminantium infection rate was less evident. There was no parallel between the prevalence of heartwater on the farms and the immune status of the animals. In general, higher tick counts were recorded in herds where strategic tick control is practised than on farms with a total tick control programme. The method of tick control did not, however, appear to influence the immune status of the cattle, the tick infection rate, or the prevalence of heartwater.     

The clinical pathology of heartwater. II. Studies on cardiac and pulmonary function in 4 calves with experimentally-induced heartwater

Studies to evaluate cardiac and pulmonary function were undertaken in 4 calves suffering from experimentally-induced heartwater. There was a marked variation in the course of the disease. Three of the calves recovered spontaneously after developing clinical signs. These included a rectal temperature in excess of 40 degrees C, anorexia and listlessness but no neurological signs. The remaining calf died 2 days after developing a fever and neurological signs. In the 3 calves that recovered, a mild hypoxemia developed during the acute stage of the disease. Arterial CO2 tension remained within normal limits, but there was a tendency towards an alkalosis. Increases in pulmonary dead space and fluctuations in venous admixture were observed. The calf that died showed similar mild changes in blood gas parameters, despite the presence of a marked reduction in minute volume, and a lung oedema was demonstrated on post-mortem examination. No marked changes in systolic and diastolic blood pressures and in right cardiac intraventricular pressures were observed. Terminally, however, there were marked decreases in stroke volume and cardiac output. These changes were associated with a sharp increase in heart rate. No primary cardiac pathology was observed on clinical and post-mortem examinations.     

The clinical pathology and pathophysiology of heartwater: a review

This paper reviews the available literature on the clinical pathology and pathophysiology of heartwater and makes comparisons with unpublished results obtained from a recent study in experimentally-induced heartwater in calves. The pathophysiological changes seem to center on an increased capillary permeability the result of which is reflected most noticeably in cardiac and lung function. There is a marked drop in cardiac output in severe cases and some workers have recorded a severe drop in diastolic blood pressure in the advanced stage of the disease. Changes in lung function are variable, depending on the stage of the disease, and may change from a respiratory alkalosis in the early febrile stage to a respiratory acidosis in more advanced cases. The basic cause for the increased capillary permeability is not known. The main clinical pathological changes measured include a progressive anaemia, fluctuations in total and differential white cell count, of which an eosinopenia and a lymphocytosis are the most marked, increases in total bilirubin which coincide with darkening of plasma colour, and a drop in total serum proteins mostly shown in the albumin levels.     

Adult Amblyomma hebraeum burdens and heartwater endemic stability in cattle.

Several conclusions of importance to studies on the epidemiology of heartwater were drawn from an investigation in which the numbers of adult Amblyomma hebraeum ticks, to which a closed herd of Hereford cattle were exposed over a period of 6 1/2 years, were manipulated. With a tick Cowdria ruminantium infection rate of 3-5%, an endemically stable situation was created by dipping the herd only when an average of 10 adult male and female A. hebraeum ticks were counted on 10 animals. When the average was increased to 15 during the calving period, 97% of calves acquired a tick-mediated immunity at the age of 6 months. Because only adult ticks confined to the hindquarters are counted, this procedure is recommended as a feasible and practical guideline to stock owners wishing to determine a dipping programme that would ensure endemic stability. The indirect fluorescent antibody test gave a true reflection of the infection rate through ticks in calves 3-6 months old, but not in older animals that had been re-infected more than once. This is because on one hand antibody may persist for 2 years after withdrawal from tick exposure and on the other the artificial re-infection of cattle with a tick acquired immunity is not always followed by a rise in antibody titres and may even result in seronegativity. Four cows infected and re-infected through ticks, remained immune to challenge for 2 years after withdrawal from tick exposure. Within the confines of one farm 3 isolates of C. ruminantium that differed in pathogenicity and immunogenicity were recovered from ticks. One of these isolates was almost non-pathogenic to cattle.     

The control of heartwater by means of tick control

Two main methods are currently being used by farmers in the heartwater regions to control this disease in cattle and other stock, namely strategic control of ticks and total control of ticks. Some farmers make successful use of the latter method, particularly in areas which are marginal for the disease. Total tick control requires top managerial skills, which includes continual expert supervision, a relatively rare asset. The majority of farmers, often unintentionally, apply strategic control of ticks to control heartwater, either without or with vaccination against the disease. In this case the objective of tick control is to minimize "tick worry" without interfering too drastically with natural transmission of the disease. This method would make provision for natural immunization of young animals (a more uncertain method than vaccination) and the maintenance of immunity. In practice these methods appear to diminish deaths from heartwater appreciably, but do not protect all animals from the disease. Furthermore, they appear to be more effective in cattle than other stock. Although strategic control has an evolutionary background and considerable merit under African conditions, it is being applied in an empirical way because of paucity of information on the epidemiology of heartwater. More information on the infection rate in and infectivity of ticks (percentage of infected ticks) offer challenging research possibilities.     

The effect of dexamethasone-induced immunosuppression on the development of faecal antibody and recovery from and resistance to rotavirus infection.

Rotavirus-naive and rotavirus-immune gnotobiotic calves were treated with high doses of dexamethasone (DX) to suppress the immune system. Calves were then infected with a virulent rotavirus inoculum, J-160, to investigate the role of immune responses both in recovery from primary rotavirus infection and in resistance to secondary rotavirus infection. Treatment of calves with DX markedly suppressed in vitro responsiveness of peripheral blood lymphocytes to mitogens within 48 h of the start of DX treatment. Suppression was similar in rotavirus-naive and rotavirus-immune calves. In contrast, the effect of DX treatment on specific antibody responses differed depending on when DX treatment started in relation to rotavirus infection. When DX treatment commenced prior to primary rotavirus infection both systemic and local specific antibody responses were inhibited. These calves, in which mitogen and antibody responses were suppressed, exhibited greater clinical signs than did control calves after infection with virulent rotavirus, but virus excretion was affected in only one of the two calves. When DX treatment was started after primary rotavirus infection but before secondary infection, systemic and local antibody responses to the primary infection and to the challenge infection were not affected. These calves resisted challenge with virulent virus as did DX-untreated rotavirus-immune calves, even though mitogen responses were suppressed. We conclude that in a primary rotavirus infection, virus excretion ceased when both antibody and mitogen responses were suppressed. Resistance to secondary rotavirus infection occurred when mitogen responsiveness was suppressed, but when antibody levels were normal. Thus, no evidence was obtained that fully functional cell-mediated immune mechanisms are essential for resistance to rotavirus infection. Evidence was provided for the ability of parenteral treatment with DX to suppress mucosal as well as systemic antibody responses.     

Genetic resistance of Guadeloupe native goats to heartwater

The resistance rate of different populations of the same breed of creole Guadeloupean goats to Cowdria ruminantium infection varied greatly depending on the previous heartwater history of each population. After experimental infection of goats removed decades ago from endemic areas, the observed resistance rate was 25%, while it was 54% in a population that had been isolated from the disease for 10 years and reached 78% in a flock actually exposed to heartwater. This resistance seems to be under genetic control as sex and paternity were the most important factors which could explain resistance in a group of 90 kids of the same flock, tested under controlled conditions. Resistance rate varied greatly (20-83%) depending on the sire, with a heritability estimate of 0.49 for half sibs and 0.85 for full sibs. A recessive sex-linked gene could be involved in the genetic determination of this resistance. From these observations, it can be stated that in endemic heartwater areas, each population, i.e. each flock, will have developed resistance at a definite rate according to population, age and the extent of past and present exposure to the disease, through a natural selection of resistant lines. Populations removed from exposure to heartwater will progressively lose their ability to resist infection through an increase in the frequency of susceptible stock. If our hypothesis of a recessive sex-linked gene is proved correct, it should be easy to select for improved resistance to the Guadeloupe breed of goat to heartwater.     

Chemo-immunization of ruminants againstBabesia,Anaplasma, Theileria andCowdria species

Current approaches to the procedure and clinical use of chemoimmunization to control babesiosis, anaplasmosis, theileriosis and heartwater in ruminants are discussed. The procedures comprise artificial infection of susceptible animals with more or less virulent strains of the causative parasites and subsequent or simultaneous treatment with an appropriate drug to moderate the reactions to the parasites. It is proposed that chemo-immunization be routinely carried out in the tropics as a method of immunizing susceptible cattle against these haemotropic infections before exposure to natural infection in the field. The method is particularly suitable for introduction of susceptible cattle into an enzootic area alongside an adequate tick control programme.     

Elaphostrongylus cervi infection in moose (Alces alces). Prevalence and pathological changes in relation to age and season

The prevalence of Elaphostrongylus cervi infection in 8–10-month-old moose calves shot in March/April in Southeastern Norway was studied. The prevalence was compared with the prevalence of infection in 4–6- and 16–18-month-old animals shot in September/October. The location of adult worms, the pathological changes which they caused within the central nervous system and their pathogenicity, were also studied. Examination of 54 8–10-month-old moose calves revealed that 46 (85 %) of the calves were infected with E. cervi. Thus a considerable proportion of the calves in the areas investigated obviously became infected with E. cervi during their first summer season. The prevalence of infection seemed to decline from the age of 8–10 months in spring till the age of 16–18 months in autumn. No difference in mean carcass weight was found between infected and non-infected calves of comparable age. Within the central nervous system adult E. cervi and associated pathological changes were found exclusively in the epidural space. The high prevalence of E. cervi infection, the insignificant differences in general condition between infected and non-infected calves and the epidural location of adult worms suggest, when considered together, that E. cervi infection in general is only moderately pathogenic in moose.The lungworm Varestrongylus alces was found in 18 of 70 moose calves examined (26 %).     

The impact of pestivirus on an artificial breeding program for cattle

After the introduction of pestivirus into a herd undergoing an embryo transfer and artificial insemination program, substantial post-weaning calf losses occurred. The predominant clinical feature was severe respiratory disease, in contrast to the commonly recognised mucosal disease. Thirty-one of 76 calves were affected, with a case fatality rate of 58%. All calves which were persistently infected with pestivirus died during the study period. There was a significant association in the surviving calves between the occurrence of recent pestivirus infection and respiratory disease. The losses on this property clearly indicate the need to routinely screen animals in an artificial breeding program for freedom from pestivirus infection.