Theileria annulata infections

Pictures

Picture	Title	Caption	Copyright
Title Two-host tick life history of Theileria infection Caption The role of Hyalomma ticks in Theleria annulata infection in cattle. Copyright Modified with permission of Elsevier Science	Two-host tick life history of Theileria infection	The role of Hyalomma ticks in Theleria annulata infection in cattle.	Modified with permission of Elsevier Science
Title Epidemiology Caption High-grade taurine calves in an endemic area of tropical theileriosis in Turkey, housed in a cattle shed which also harbours hyalommid ticks. Copyright The University of Edinburgh	Epidemiology	High-grade taurine calves in an endemic area of tropical theileriosis in Turkey, housed in a cattle shed which also harbours hyalommid ticks.	The University of Edinburgh
Title Epidemiology Caption Cattle-shed in an endemic area of tropical theileriosis in the Black Sea Coastal area of Turkey. Copyright The University of Edinburgh	Epidemiology	Cattle-shed in an endemic area of tropical theileriosis in the Black Sea Coastal area of Turkey.	The University of Edinburgh
Title Cytokine production by Theileria annulata Caption Cytokine production by Theileria annulata schizont-infected cells in lymph node of calf undergoing tropical theileriosis: merozoite-infected cells (red) labelled with an antibody to TNF-alpha and the avidin-biotin complex/alkaline phosphatase (ABC/AP) system (Original x 1000). Copyright Used with permision from Academic Press Ltd.	Cytokine production by Theileria annulata	Cytokine production by Theileria annulata schizont-infected cells in lymph node of calf undergoing tropical theileriosis: merozoite-infected cells (red) labelled with an antibody to TNF-alpha and the avidin-biotin complex/alkaline phosphatase (ABC/AP) system (Original x 1000).	Used with permision from Academic Press Ltd.
Title Theileria annulata disease course Caption Eye of cow infected with T. annulata showing anaemic mucous membranes with characteristic haemorrhaging. Copyright The University of Edinburgh	Theileria annulata disease course	Eye of cow infected with T. annulata showing anaemic mucous membranes with characteristic haemorrhaging.	The University of Edinburgh
Title Disease course Caption Haemorrhages in the skin around mammary glands of cow infected with T. annulata. Copyright The University of Edinburgh	Disease course	Haemorrhages in the skin around mammary glands of cow infected with T. annulata.	The University of Edinburgh
Title Carcase showing the advanced stages of Tropical Theileriosis Caption Haemorrhages on mesenteric lymph nodes. Copyright The University of Edinburgh	Carcase showing the advanced stages of Tropical Theileriosis	Haemorrhages on mesenteric lymph nodes.	The University of Edinburgh
Title Cow with Theileriosis (Theileria annulata) Caption Taurine cow infected with T. annulata showing debilitated condition. Copyright The University of Edinburgh	Cow with Theileriosis (Theileria annulata)	Taurine cow infected with T. annulata showing debilitated condition.	The University of Edinburgh
Title Carcase showing the advanced stages of Tropical Theileriosis Caption Yellow connective tissue and haemorrhages on body wall Copyright The University of Edinburgh	Carcase showing the advanced stages of Tropical Theileriosis	Yellow connective tissue and haemorrhages on body wall	The University of Edinburgh
Title Theileria annulata infections. Advanced stage of Tropical Theileriosis. Caption Cortical haemorrhages in the kidney. Copyright The University of Edinburgh	Theileria annulata infections. Advanced stage of Tropical Theileriosis.	Cortical haemorrhages in the kidney.	The University of Edinburgh
Title Theileria annulata infections. Parasitized mononuclear cells in kidney. Caption In tissues of calves undergoing tropical theileriosis: schizont-infected cells in the glomerulus of the kidney during late stages of the disease. Schizonts (brown) labelled with the ABC/HRP system (brownish colour) (Original x 1000) Copyright Used with permission from Academic Press Ltd.	Theileria annulata infections. Parasitized mononuclear cells in kidney.	In tissues of calves undergoing tropical theileriosis: schizont-infected cells in the glomerulus of the kidney during late stages of the disease. Schizonts (brown) labelled with the ABC/HRP system (brownish colour) (Original x 1000)	Used with permission from Academic Press Ltd.
Title Advanced stage of Tropical Theileriosis Caption Grossly enlarged and haemorhagic lymph node draining the site of inoculation of T. annulata sporozoites. Copyright The University of Edinburgh	Advanced stage of Tropical Theileriosis	Grossly enlarged and haemorhagic lymph node draining the site of inoculation of T. annulata sporozoites.	The University of Edinburgh
Title Advanced stage of Tropical Theileriosis Caption Echymotic haemorrhages on mesenteric lymph node. Copyright The University of Edinburgh	Advanced stage of Tropical Theileriosis	Echymotic haemorrhages on mesenteric lymph node.	The University of Edinburgh
Title Advanced stage of Tropical theileriosis Caption Haemorrhages in abomasal wall. Copyright The University of Edinburgh	Advanced stage of Tropical theileriosis	Haemorrhages in abomasal wall.	The University of Edinburgh
Title Advanced stage of Tropical Theileriosis Caption Abomasal ulcers. Copyright The University of Edinburgh	Advanced stage of Tropical Theileriosis	Abomasal ulcers.	The University of Edinburgh
Title Parasitized mononuclear cell Caption In tissues of calves undergoing tropical theileriosis: schizont-infected cells (brown) in the paracortex of the prescapular lymph node draining the site of inoculation of a calf undergoing the middle stage of disease. Schizonts labelled with an anti-schizont monoclonal antibody and the avidin-biotin complex/horse radish peroxidase (ABC/HRP) system x 1000 Copyright Used with permission from Academic Press Ltd.	Parasitized mononuclear cell	In tissues of calves undergoing tropical theileriosis: schizont-infected cells (brown) in the paracortex of the prescapular lymph node draining the site of inoculation of a calf undergoing the middle stage of disease. Schizonts labelled with an anti-schizont monoclonal antibody and the avidin-biotin complex/horse radish peroxidase (ABC/HRP) system x 1000	Used with permission from Academic Press Ltd.
Title Parasitized mononuclear cells Caption In tissues of calves undergoing tropical theileriosis: small numbers of schizont-infected cells (brown) and many merozoite-infected cells (stained blue with Harris's haematoxylin) in the medullary sinuses of the lymph nodes during the late stages of disease (Original x 1000) Copyright Used with permission from Academic Press Ltd.	Parasitized mononuclear cells	In tissues of calves undergoing tropical theileriosis: small numbers of schizont-infected cells (brown) and many merozoite-infected cells (stained blue with Harris's haematoxylin) in the medullary sinuses of the lymph nodes during the late stages of disease (Original x 1000)	Used with permission from Academic Press Ltd.
Title Parasitized mononuclear cells Caption In tissues of calves undergoing tropical theileriosis: large numbers in the wall of the abomasum. Schizonts (brown) labelled with ABC/HRP system (Original x 100) Copyright Used with permission from Academic Press Ltd.	Parasitized mononuclear cells	In tissues of calves undergoing tropical theileriosis: large numbers in the wall of the abomasum. Schizonts (brown) labelled with ABC/HRP system (Original x 100)	Used with permission from Academic Press Ltd.
Title Parasitized mononuclear cells in the spleen Caption In tissues of calves undergoing tropical theileriosis: schizont-infected cells in the white and red pulp of the spleen during late stage of disease. Schizonts (brown) labelled with ABC/HRP system (Original x 100) Copyright Used with permission from Academic Press Ltd.	Parasitized mononuclear cells in the spleen	In tissues of calves undergoing tropical theileriosis: schizont-infected cells in the white and red pulp of the spleen during late stage of disease. Schizonts (brown) labelled with ABC/HRP system (Original x 100)	Used with permission from Academic Press Ltd.
Title Histopathology Caption Histiocytic response in paracortex of draining lymph node during advanced stage of tropical theileriosis, stained with Haematoxylin and Eosin (Original x 100). Copyright The University of Edinburgh	Histopathology	Histiocytic response in paracortex of draining lymph node during advanced stage of tropical theileriosis, stained with Haematoxylin and Eosin (Original x 100).	The University of Edinburgh
Title Diagnosis Caption Smear of biopsy material from lymph node draining the site of inoculation showing T. annulata schizont-infected cells. Copyright The University of Edinburgh	Diagnosis	Smear of biopsy material from lymph node draining the site of inoculation showing T. annulata schizont-infected cells.	The University of Edinburgh
Title Diagnosis Caption Intra-erythrocytic piroplasms of Theileria annulata: typical ring forms in a smear made from the blood of a calf undergoing the advanced stages of tropical theileriosis. Copyright Used with permission from CABI Publishing	Diagnosis	Intra-erythrocytic piroplasms of Theileria annulata: typical ring forms in a smear made from the blood of a calf undergoing the advanced stages of tropical theileriosis.	Used with permission from CABI Publishing
Title Diagnosis Caption Antibodies to T. annulata schizonts detected in an immunofluorescent antibody test (IFAT) using T. annulata schizont-infected cells as antigen and a fluorescein labelled conjugate to detect the reaction between the antibodies and the antigen. Copyright The University of Edinburgh	Diagnosis	Antibodies to T. annulata schizonts detected in an immunofluorescent antibody test (IFAT) using T. annulata schizont-infected cells as antigen and a fluorescein labelled conjugate to detect the reaction between the antibodies and the antigen.	The University of Edinburgh
Title Vaccines Caption T. annulata schizont-infected cells growing in tissue culture. After prolonged culturing, such cultures become attenuated and can be used as vaccines. Copyright The University of Edinburgh	Vaccines	T. annulata schizont-infected cells growing in tissue culture. After prolonged culturing, such cultures become attenuated and can be used as vaccines.	The University of Edinburgh

Identity

Preferred Scientific Name

• Theileria annulata infections

International Common Names

• English: Egyptian fever; Mediterranean Coast fever; theileriosis in ruminants - exotic; theileriosis, tropical; tropical theileriosis
• French: theileriose bovine d'Afrique du Nord
• Chinese: huan xing tai le chong bing

Local Common Names

• India: chichri rog
• Tunisia: boussofie

Pathogen/s

Overview

Theileria annulata is a tickborne intracellular protozoan parasite of cattle, which occurs in North Africa, southern Europe, the Near and Middle East, India, China and Central Asia. It causes both mortality and reduced production, and has significant economic impacts as a result.

Datasheets are also available on bovine theilerioses and theileriosis in general (as well as other specific Theilieria infections).

Host Animals

Systems Affected

Distribution

Updated information on the distribution of theileriosis in general (as distinct from Theileria annulata infections in particular) can be found in OIE's WAHID database on disease occurrence: http://www.oie.int/en/links/wahid/.

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Pathology

The chief pathological features of acute lethal tropical theileriosis are associated with dysfunction of the lymphoid and reticulo-endothelial system, and haematological changes (Dschunkowsky and Luhs, 1904; Neitz, 1957; Pipano, 1974, 1994; Levine, 1985). The general post-mortem findings (Figures 35-41) are as follows:
Carcasses: emaciated; anaemic; icteric; yellowish and gelatinous connective tissues.
Blood: appears watery.
Subcutaneous tissues: numerous petechial or larger haemorrhages.
Mucous membranes (pharynx, larynx, trachea, bronchi): pale; numerous petechial or haemorrhages.
Serous membranes: pale with numerous petechial or larger haemorrhages.
Lymph nodes: are markedly enlarged, oedematous, have varying degrees of haemorrhage, are frequently hyperplastic and hyperaemic.
Thymus: mildly congested.
Heart: myocardial muscular degeneration, petechial and ecchymotic haemorrhages of the epi- and endocardium.
Lungs: frequently oedematous and congested.
Spleen: splenomegaly due to lymphoid hyperplasia, spleen is soft and may have prominent malpighian corpuscles.
Liver: hepatomegaly; pale brown or yellow, friable, with evident parenchymatous degeneration.
Gall bladder: often markedly distended, produces dark-green viscid bile.
Kidneys: pale and congested, pseudoinfarcts of lymphoproliferative foci.
Adrenal glands: cortical haemorrhages, severely infiltrated.
Abomasum, small and large intestines: characteristic ulcers surrounded by a haemorrhagic zone.
Nervous system: occasionally cerebral haemorrhages (Sharma and Gautam, 1973).
Bladder: dark-brown urine due to bile pigments.
Skin: exceptionally non-pigmented papules or pustules, necrotic, ulcerative (Manickam et al., 1984).
Eye: exopthalmos observed in cattle in Iran (Baharsefat et al., 1977), exophthalmos and ulceration of the conjunctiva observed in Iraq (Khalifa and Kadhim, 1967), ocular lesions in India (Mallick et al., 1986; Aher et al., 1990).

Specific post-mortem findings have been reported for cases in Bulgaria (Pavlov, 1957; Vulhovski and Pavlov, 1970), Trans-Caucasia (Dschunkowsky and Luhs, 1904), Central Asia (Oboldoueff and Galuzo, 1928), Algeria (Sergent et al., 1945), India (Sen and Srinivasan, 1936; Prasad, 1946; Gautam et al., 1970; Sharma and Gautam, 1971; Gill et al., 1977; Srivastava and Sharma, 1981); Iran (Rafyi and Maghami, 1962; Baharsefat et al., 1977), Iraq (MacHattie, 1935; Khalifa and Kadhim, 1967).

Diagnosis

For information on diagnosis, see the 'bovine theilerioses' datasheet. The  indirect fluorescent antibody test (IFAT) is a prescribed test for international trade, and is described in OIE's Manual of Diagnostic Tests and Vaccines for Terrestrial Animals (OIE, 2013; http://www.oie.int/fileadmin/Home/eng/Health_standards/tahm/2.04.16_THEILIERIOSIS.pdf).

List of Symptoms/Signs

Disease Course

Tropical theileriosis or Mediterranean Coast fever is a non-contagious disease of cattle characterized by fever, enlarged peripheral lymph nodes, emaciation, progressive anaemia and jaundice (Brown, 1990a).T. annulata causes similar disease symptoms across most of its range, as reported in Central Asia (Oboldoueff and Galouzo, 1928; Markov, 1962), in North Africa (Sergent et al., 1945), in the Middle East (Pipano, 1974; El-Metenawy, 2000), in China (Luo and Lu, 1997), in Iran (Rafyi and Maghami, 1962) and in India (Sen and Srinivasa, 1936; Gautam et al., 1970; Sharma and Gautam, 1971; Prasad, 1946). These similarities exist in spite of T. annulata exhibiting extensive strain variation in terms of virulence (Sergent et al., 1945; Pipano, 1974) and molecular diversity (Ben Miled et al., 1994; Katzer et al., 1998; Gubbels et al., 2000). Animals may survive disease but recovery and convalescence can be prolonged and incomplete, leading to permanent debilitation, loss of productivity and a prolonged carrier state (Uilenberg, 1981). Milk production is depressed and may not re-establish itself entirely, even after the following calving (Markov, 1962). In the Asian buffalo, which is probably its original host, T. annulata causes only subclinical or very mild infections (Hadier, 1992; Uilenberg, 1995).

T. annulata causes an acute, mild, sub-acute, per-acute or chronic disease, according to strain virulence and host susceptibility (Sergent et al., 1945; Neitz, 1957; Pipano, 1994; Luo and Lu, 1994). In the acute lethal disease, cattle exhibit increasingly pronounced symptoms from five to seven days after infection, when schizonts become detectable in the lymph node draining the site of infection (Preston et al., 1992).

The early clinical signs include pyrexia coinciding with the appearance of schizonts in the lymph node draining the site of inoculation, and also leucopenia, enlarged superficial lymph nodes, inappetance, drooling, serous nasal discharge, swelling of the eyelids, drooping ears, lowered head, sluggish gait, lacrimation, accelerated pulse, general weakness, decreased milk production and sometimes nervous symptoms. As the disease progresses, the following signs occur: cachexia followed by wasting, marked anaemia with bilirubinaemia and bilirubinurea, superficial lymph nodes become greatly enlarged, thrombocytopenia, conjunctiva icteric with petechial haemorrhages and diarrhoea. During the later stages, the faeces are often mixed with blood and mucus, the skin may bear petechiae and, more rarely, raised nodules containing schizont-infected cells. There may be marked emaciation and recumbency. If erythrocytes do not regenerate, anaemia becomes so severe, and dyspnoea so pronounced, that death ensues 8-15 days after the onset of the disease.

Death may occur within two to three weeks following the proliferation of schizont-infected cells or the haemolytic anaemia resulting from intra-erythrocytic piroplasms. Parasitaemias may reach 60% during the terminal stages of the disease and more rarely, following acute disease, up to 90%. Very rarely, acute infections are accompanied by cerebral symptoms (Sharma and Gautam, 1973; Srivastava and Sharma, 1976). Per-acute disease is fairly common and onset is sudden. Animals with per-acute disease exhibit the symptoms of acute disease, but death preceded by hypothermia occurs within a few days. A panleucopaenia and thrombocytopaenia are common features of acute and per-acute disease syndromes (Prasad, 1946; Laiblin, 1978; Brown, 1990a; Preston et al., 1992). Mild strains may cause subclinical infections, with mild symptoms lasting a few days or sub-acute disease with an irregularly intermittent fever lasting for two to four weeks and less marked symptoms than in acute infections. Animals usually recover but pregnant animals may abort. Chronic disease may persist for a month or more with animals taking more than two months to recover. Sub-acute, acute or chronic forms of the disease sometimes lead to acute episodes and death. Animals dying after a prolonged course show evidence of having suffered a severe aplastic anaemia (Laiblin, 1978; Pipano, 1994). There is evidence for neonatal infections with T.annulata (Mishra et al., 1994).

Anaemia in T. annulata infections has been linked to parasitaemia (Preston et al., 1992) exacerbated by the proliferation of intra-erythrocytic stages (Conrad et al., 1985), autoimmune mechanisms (Hooshmand-Rad, 1976) and erythro-phagocytosis by macrophages (Forsyth et al., 1999). Extensive removal of erythrocytes causes bilirubinaemia and biliruburea in T. annulata infections (Barnett, 1977).

Protective immunity to T. annulata infections

The different parasite stages of T. annulata appear susceptible to attack by a variety of innate and adaptive immune responses (Preston et al., 1999). Parasitized macrophages may be lyzed by CD8+ T-cells and Natural Killer (NK) cells. Macrophage-derived nitric oxide may eliminate sporozoites, trophozoites and schizont-infected cells. Parasite-inhibitory cytokines (e.g. IFN-g ) may kill sporozoites and prevent establishment of trophozoite-infected cells.

The immunobiology of T. annulata and T. parva is reviewed by Morrison (2009), and immune responses to T. annulata by Ahmed et al. (2008).

Epidemiology

The hyalommid vectors of T. annulata occur in sub-humid and semi-arid environments, semi-desert and desert habitats, steppes and dry-grass lands (Galouzo et al., 1958). The most important vectors are Hyalomma anatolicum anatolicum, a three-host tick, and Hyalomma detritum, a two-host tick (Barnett, 1968, 1977; Uilenberg, 1981). These species can act as vectors because all stages have adapted to cattle (Jongejan and Uilenberg, 1994). H. scupense may serve as a field vector (Stepanova, 1976; Markov, 1958; Galouzo et al., 1958), when animals live in close contact (Markov, 1962). Few of the two-host ticks that transmit T. annulata experimentally are likely to be field vectors, because only the adults feed on cattle (Hadani et al., 1963; Jongejan et al., 1983; Um El Hassan et al., 1983). Reports of hereditary transmission(Markov, 1962; Ray, 1950) remain unsubstantiated.

Impact: Economic

A global figure for the economic impact and importance of T. annulata is not available, but this parasite is accepted to affect the productivity of perhaps 250 million cattle, causing morbidity in indigenous cattle and mortality of between 40-60% 'improved' cross-bred or exotic animals (Brown, 1990b). It is probably the most important of all tick-borne diseases of domestic stock (Purnell, 1978). The economic impact of T. annulata in India was estimated to be US$ 800m, based on direct losses due to mortality and production losses (milk yield, growth rate, meat, infertility, abortion, calving interval and hides) and the indirect costs of control measures (dipping, vaccination, chemotherapy, veterinary legislation and monitoring) (Brown, 1997). Estimates for Gujarat put the cost of infections at US$ 1.27million for 40,000 cows, with a projected total loss of US$ 210 million per annum for 6.6 million adult cross-bred cows in India (Singh, 1991b). Losses attributable to T. annulata in Tunis were estimated to be approximately US$ 24 per dairy cow, taking into account mortality, treatment costs, milk yield and abortion, and including the effects of subclinical infection as well as overt disease (Darghouth, 1999).

Disease Treatment

The napthoquinones parvaquone (Hawa et al., 1988; Gill et al., 1981; McHardy et al., 1983) and buparvaquone (McHardy et al., 1985; McHardy, 1989; McHardy, 1991), and the febrifuginone halofuginone lactate (Schein and Voigt, 1979, 1981), will cure clinical disease resulting from infection with T. annulata or T. parva. However, the therapeutic dose of halofuginone (1.2 mg/kg) often produces side effects (Schein and Voigt, 1979). Parvaquone does not effect a parasitological cure and recovered animals may take several months to return to a normal level of productivity (Dolan, 1986).

Buparvaquone is a safe and effective drug, which can be used both prophylactically (during the prepatent/incubation period) and therapeutically (during patent disease) against T. annulata (McHardy et al., 1985; Dhar et al., 1987, 1988, 1990; McHardy, 1991; Sharma and Mishra, 1990; Singh et al., 1993) (as well as T. parva: McHardy et al., 1985; McHardy, 1991; Dolan et al., 1992). However, in young calves, haematopoiesis-stimulating drugs must be applied together with buparvaquone to avoid the deleterious effects of severe anaemia (Dhar et al., 1988).

Tetracyclines are effective against the schizonts of T. annulata (Gill et al., 1978; Jagdish et al., 1979; Pipano et al., 1981; Mallick et al., 1987) and of T. parva (Dolan, 1981), but only when used in large doses during the prepatent/incubation period of infection (Hashemi-Fershaki and Shad-Del, 1974).

The 8-aminoquinolones - pamaquin and primaquine - are active against the piroplasms of T. annulata (Zhang, 1987, 1997; Luo and Lu, 1997). Extracts of the plant Perganum harmala have a marked suppressive effect on natural infections of T. annulata (Hu et al., 1997).

Prevention and Control

Immunization is the chief weapon in the control of tropical theileriosis in highly susceptible dairy breeds of cattle. Attenuated schizont-infected cell culture vaccines remain the main type of vaccine in use (OIE, 2013), notwithstanding progress towards the development of subunit vaccines (Morrison and McKeever, 2006).

The development and application of the attenuated vaccines has been comprehensively described (Pipano, 1989a, b, 1995; Stepanova and Zablostskii, 1989). Immunization may confer protection for periods from a year (Hashemi-Fesharki, 1998) to more than three years (Zablotsky, 1991). The duration of protection in the field depends upon the existence of and level of natural tick-derived sporozoite challenge (Beniwal et al., 2000). Detailed methods for culturing Theileria schizont-infected cells and producing vaccine cell lines have been described by Brown (1983, 1987). The use of vaccination against tick-borne diseases in the development of control strategies in relation to the three main husbandry systems, cattle at pasture, cattle in barns or cattle at zero-grazing, was described by Pipano and Grewal (1990).

Countries where vaccines modelled on the Israeli cell culture vaccine have been developed and used in the field include: Iran (Hashemi-Fesharki and Shad-del, 1973; Hashemi-Fesharki, 1988, 1991, 1998), the former USSR (Stepanova et al., 1976, 1982; Stepanova, 1983; Stepanova et al., 1986; Stepanova and Zablotsky, 1989), India (Singh, 1991a, 1990; Singh et al., 1993a), China (Gansu Provincial Institute of Veterinary Medicine, 1975; Zhang, 1991; Lu and Yin, 1994; Lu and Luo, 1997; Guo et al., 1997; Song, 1997), Turkey (Ozkoc and Pipano, 1981; Ozkoc et al., 1989) and Morocco (Ouhelli et al., 1980). The efficacy of such vaccines in protecting enormous numbers of cattle over very large areas has been demonstrated by the very successful vaccination campaigns in China (Lu and Yin, 1994; Lu and Luo, 1997; Guo et al., 1997; Song, 1997).

In spite of the cross-immunity which exists between stocks of T. annulata throughout its range (Rafyi et al., 1965; Preston and Brown, 1988), the potential for transferring other pathogens has prevented the global use of one vaccine line. Therefore, countries requiring vaccines, for example Tunisia (Darghouth et al., 1996c) or Spain (Viseras et al., 1997; Viseras and Garcia Fernandez, 1999), or in some cases regions within larger countries such as China (Guo et al., 1997; Song, 1997), have had to develop their own individual attenuated vaccines. Aspects that have been studied as part of this include:

•  Identification of target populations (Darghouth et al., 1997)
   
• The desirability, or otherwise, of the carrier status
   
• The duration of immunity needed
   
• The length of time between boosters (Darghouth, 1999)
   
• The problems of suitability of vaccines for immunizing very young calves and pregnant cattle
   
• The problem of rejection when the same cell line is used to boost as well as immunize (Beniwal et al., 1997; Grewal et al., 1997; Nichani et al., 1997).

Mechanisms underlying attenuation have been studied with the aim of finding ways of improving/accelerating the attenuation process (Tait and Hall, 1990; Sutherland et al., 1996; Hall et al., 1999; Adamson et al., 2000; Boulter and Hall, 2000).

The infection and treatment method has been investigated with reference to overcoming the problem of vaccinating young calves with a cell-line vaccine (Gill et al., 1976). The drug of choice for this may be buparvaquone (Dhar et al., 1990; McHardy, 1991).

Darghouth (2008) reviews experience with live attenuated vaccines against T. annulata in Tunisia.

Practical problems associated with preparation, storage, transport, administration, possibility of transferring other pathogens and reverting to virulence, are driving work on developing sub-unit vaccines against T. annulata (Tait and Hall, 1990; Preston et al., 1999; Boulter and Hall, 2000; Hall et al., 2000). These include investigations on genetic diversity (Katzer et al., 1998; Gubbels et al., 2000; Morrison and McKeever, 2006).

Some further information on prevention and control, including vector control, breeding of resistant cattle and combinations of different control methods, is provided in the 'bovine theilerioses' datasheet.

References

Abd-El-Salam MN; Ali HS; Sadiek AH; Zaitoun AM; Sayed AS; Abu-El-Eneen GS; Abd-El-Salam FA, 1994. Haematological picture and serum protein analysis in fattening buffalo calves naturally infested with Theileria annulata. Assiut Veterinary Medical Journal, 30(60):186-193; 19 ref.

Adamson RE et al., 2000. Theileria annulata: reduced host MMP activity profile of long-term culture is transmitted as a parasite-encoded trait. Experimental Parasitology, 94:45-47.

Aher VD; Bhikane AU; Gaikwad BB; Singh B, 1990. A note on ocular lesions associated with tropical theileriosis (Theileria annulata) infection in calves. Journal of Bombay Veterinary College, 2(2):123-124; 4 ref.

Ahmed BA; Al Hoty KF; Saddik MM, 1987. Theileria annulata infection in Friesian cattle imported to Kuwait from W. Germany. Veterinary Medical Journal, Giza, Egypt, 35(1):139-146; 16 ref.

Ahmed JS; Glass EJ; Salih DA; Seitzer U, 2008. Innate immunity to tropical theileriosis. Innate Immunity, 14(1):5-12. http://ini.sagepub.com/cgi/reprint/14/1/5

Anandan R; Ganesamoorthy M; John L; Kandavel E; Lalitha CM, 1983. An account of theileriosis among calves. Cheiron, 12(6):327-328; 9 ref.

Anandan R; Lalitha CM, 1984. Bovine tropical theileriosis in Tamil Nadu. Cheiron, 13(1):34-39; 18 ref.

Angelovski T; Iliev A, 1977. Piroplasmosis and their vectors in cattle, sheep and horses of Serbian Macedonia. The First Mediterranean Conference on Parasitology, 5-11th 0ctober 1977, Izmir, Turkey. 22.

Arshadi M, 1976. Control of tick-borne diseases in Iran (Anaplasmosis, piroplasmosis and theileriosis). Bulletin de l'Office International des Epizooties, 86:45-53.

Arslan SH; Shukur AA, 1994. Clinical and haematological studies on theileriosis and anaplasmosis in local breed cattle in Mosul region. Iraqi Journal of Veterinary Sciences, 7(2):93-100; 23 ref.

Askarov EM, 1975. Virulence and immunogenicity of Theileria annulata. Veterinariya-Moscow, 5:76-78.

Badalov ET, 1989. Preventing piroplasmosis in cattle [in Tadzhikistan]. Veterinariya (Moskva), No. 6:37-40.

Baharsefat M et al., 1977. Unusual cases of Theileria annulata infection in calves. Archives de l'Insitute Razi, 29:47-58.

Bahri S; Kallel A; Gouia A, 1995. La theileriose bovine en Tunisie, etude retrospective sur 5 ans. Bulletin Epidemiologique et d'Information Veterinaire, Institut de la Recherche Veterinaire de Tunis, 5:1-3.

Barnett S, 1960. Connective tissue reactions in acute fatal East Coast fever (Theileria parva) of cattle. Journal of Infectious Diseases, 107:253-282.

Barnett SF, 1977. Theileria. In: Kreier JP, ed. Parasitic Protozoa, Vol VI. New York, USA: Academic Press, 77-113.

Ben Miled L, 1994. Phenotypic and molecular analysis of isolates of Theileria annulata. Archives de l'Institut Pasteur de Tunis, 71(3-4):463-464.

Beniwal RK; Nichani AK; Sharma RD; Rakha NK; Suri D; Sarup S, 1997. Responses in animals vaccinated with the Theileria annulata (Hisar) cell culture vaccine. Tropical Animal Health and Production, 29(4 (supplement)):109S-113S; 7 ref.

Beniwal RK; Sharma RD; Nichani AK, 2000. Determination of duration of immunity of calves vaccinated with the Theileria annulata schizont cell culture vaccine. Veterinary Parasitology, 90(1/2):25-35; 24 ref.

Bondopadhyay LN; Maity B; Bhandari SK; Gupta R, 1992. Prevalence of bovine theileriasis in and around Bethuadahari district - Nadia, West Bengal. Indian Journal of Animal Health, 31(1):79-80; 3 ref.

Boulter N; Hall R, 1999. Immunity and vaccine development in the bovine theilerioses. Advances in Parasitology, 44:41-97; 14 pp. of ref.

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