Haemorrhagic septicaemia vaccines

PII: SO264-410X(97)00313-7

Vaccine, Vol. 16, No. 11112, pp. 1184-1192, 1998 0 1998 Elsevier Science Ltd. All rights reserved Printed in Great Britain 0264-410X/98 $19+0.00

ELSEVIER

Haemorrhagic septicaemia vaccines Rishendra

Verma*S

and T.N. Jaiswa1-f

Haemowhagic septicaemia (HS), an economically important disease of cattle and buffaloes, is caused by Pasteurella multocida (6:B). Vaccination against this disease is widely practised. Plain broth bacterins, or alum precipitated and aluminium hydroxide gel vaccines are administered twice a year since these vaccines offer an immunity of 4-6 months. Many countries use oil adjuvant vaccine (OAV), which gives both a higher degree and a longer duration of immunity up to 1 year A double emulsion and multiple emulsion vaccine consisting of a thin viscosity have also been experimentally developed that gave an immunity parallel to OAV Recently, a live vaccine developed from a fallow deer strain (B:3,4) has been used in Myanmar that offers an immunity for more than a year but is not free from constraints. The present review provides information on HS vaccines developed from time to time using whole bacteria or their components. The kinetics and isotype of antibody and cell-mediated immune responses have also been poorly understood so far and hence information on their role in protection against HS is reviewed. 0 1998 Elsevier Science Ltd. All rights reserved Keywords:

Haemorrhagic

septicaemia

vaccines;

immune

responses

Pasteurellosis in cattle and water buffalo is a specific form of acute and fatal disease commonly referred as ‘Haemorrhagic septicaemia’ (HS), which is caused by two specific serotypes of Pasteurella multocida. The Asian serotype is designated B:2, and the African serotype is E:2 by the Carter-Heddleston system, corresponding to 6:B and 6:E by the Namioka-Carter system. HS is considered economically to be the most important disease in Southeast Asia’, including Indonesia, Phillipines, Thailand, Malaysia, Middle East, North East, central and South Africa. The disease has also been reported in American Bison in 1967 and dairy cattle in 1969 in USA’. A low sporadic incidence of HS has been reported in many Southern European States and the former USSR. It was estimated a decade ago that the annual losses to the North American cattle industry due to the disease were approximately $USSOO million’. Recently, annual losses of $US400-6000 have been reported in Indonesia, $US1.4 million in Laos and approximately $USl .O million in Malaysia3. Fifty-three thousand bovine deaths in India are attributed to this disease annually5. Vaccination as a deliberate attempt to protect animals against disease has a long history, although only in the 20th century has the practice developed into routine vaccinations. Pasteur6 by chance discovered that a weakened fowl cholera (tl multocida) culture, left exposed to the air over a holiday, provided *Division of Standardisation, lzatnagar 243 122 (UP), India. tDivision of Bacteriology and Mycology, Indian Veterinary Research Institute, lzatnagar 243 122 (UP), India. SAuthor to whom all correspondence should be addressed. Fax: +91 581 457284. (Received 8 January 1997; revised version received 10 November 1997; accepted 4 December 1997)

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immunity against a challenge with virulent organisms. He was quick to notice that the principle was the same as that discovered by Jenner’, although attenuation had been achieved in a different manner. Pasteur6 observed that his vaccine would be more pure because it consisted of a weakened form of the same organism that caused the disease. The modern concept of vaccination, therefore, was really introduced by Pasteur6 with the chicken cholera vaccine. Similarly, the best way to prevent haemorrhagic septicaemia (HS) in cattle and buffaloes is vaccination. The present review deals with the development of haemorrhagic septicaemia vaccines, immunity offered by them and their role in protection.

KILLED VACCINES Plain bacterin

Baldrey’ first advocated prophylactic vaccination of cattle against HS with killed (0.25% lysol-inactivated broth) vaccine that offered immunity for 6 weeks. The protective value of vaccine was investigated by Holmes’, and heat-killed broth cultures mixed with a gluteraldehyde extract of old culture sediments produced solid imunity for 6 weeks in 70% of the vaccinated animals. Holmes’” and Baldrey” developed a sensitized vaccine by mixing washed live orgnaisms with the corresponding hyperimmune serum and resuspended the organisms in saline after allowing the mixture to mix on room temperature at 24 to 48 h. The saline suspension was subsequently heated to 60°C for 30 min. The vaccine was not eventually used due to the large ,quantities of serum needed for its manufacture. Allen thought that the dead vaccine induced protection against HS under field conditions for at least a year. Chaeh’” observed the effect of a 72-h broth bacterin against HS that was protective but was found

Haemorrhagic

to be inferior to alum-precipitated vaccine in respect of the duration of immunity. Chorherr14 studied formolinor heat-inactivated !! multocida vaccines. The index of protection was higher with the heat-inactivated vaccine than the formolized vaccine. He found no improvement in its protectivity by the addition of various adjuvants. A vaccine was prepared from a strain isolated from cattle and buffaloes that consisted of a suspension of lysed bacteria in 5% saponin to provide a continuous antigenic stimulus by retarding its absorption at the site of inoculation of the vaccine. Satisfactory results of this vaccine have been claimed in Iran where it was widely used. However, this vaccine did not find much use due to its undesirable tissue-damaging effect”,‘“. Poor results with a saponin vaccine on smallscale laboratory experiments have been reported”. However, later, the saponin lysed vaccine was replaced with saponin-formolized vaccinelx. Agar-wash

vaccine

Due to the protein shock encountered in some animals vaccinated with broth vaccine, an agar-wash using the formolin-killed cultures was vaccine developed at IVRI’“. Rau and Govil”’ developed an improved agar-wash heat-killed or formolized vaccine using a higher concentration of cells, which provided immunity for 4 months. Alum-precipitated

vaccine

Alum-precipitated vaccine was apparently tried by American commercial manufacturers prior to the 1950s” and also in the Phillipines. The vaccine has been extensively used in Southern India’” and in Central Africa. It has also been used in the former USSR and appears to be the most popular vaccine used in Asian countries. Nikiphorova-3 reported that alum-precipitated vaccine protected cattle, sheep and goat against p multocida infection for 6 months. Controlled trials in Sri Lanka using a standard vaccine containing 2.5 mg of bacteria zger dose gave 3-4 months’ immunity to challenge . To reduce the incidence of shock reactions, in Thailand, vaccine is prepared from the cultures produced in vortex aeration and blended with aluminium tanks formolized hydroxide gel (0.3%)‘5. Alum-precipitated vaccine in conjunction with levamisole has been reported to increase antibody titres significantly’“. The efficacy of vaccine in rotection against alum-precipitated P challenge for HS has been reported-‘.“. Aluminium

hydroxide gel vaccine

Thailand and Laos use an aluminium hydroxide gel vaccine extensively. Immunity up to 4 months is claimed. Further aluminium hydroxide gel HS vaccine did not confer a substantial immunity of more than 90 days even using Vitamin E and levamisole as immunomodulators”. Adjuvanted

vaccines

Simple emulsions with little stability can be formed by shaking together two immiscible liquids. However, if permanence and stability are required, a third substance, the emulsifying agent, must be introduced.

septicaemia

vaccines: R. Verma and T.N. Jaiswal

The choice of emulsifying agent is very important

in influencing the type of emulsion that results, e.g. waterin-water. Lanolin has been chosen as the emulsifying agent in HS vaccine because it is cheap, readily available, effective and non-irritating. Another agent widely used for water-in-oil emulsions is mannite monooleate, a commercial brand of which is Arlacel A. However, it has been shown that the emulsions made with mannite monooleate are no more stable than those prepared with modern lanoli?“. In Burma, as early as 1952, a small quantity of mineral oil adjuvant vaccine using a field strain ‘insein’ was prepared that provided good protection. Subsequently, a new strain ‘Katha’ isolated in Burma was found to yield superior results. Work on the development of an adjuvant vaccine in India was initiated at IVRI in 1953j’. Adjuvants enhance protection in cattle against a higher challenge dose of virulent organisms than bacteria alone”. The use of oil adjuvant-inactivated vaccine against P multocida infection in farm animals has been investigated”‘.““. A passive immunization trial in mice showed that oil-adjuvanted vaccines were more superior than a bacterial suspension when tested at 25 weeks postinoculation”‘. Iyer et al.” studied the effect of alum, mineral oil, lanoline mixture and liquid paraffinlanoline mixture as adjuvants. After large-scale field and laboratory trials over several years, yeast extract agar peptone (YEAP) grown, washed, formolized (0.2%) vaccine mixed with liquid paraffin lanolin adjuvants was found to be safe for cattle and buffaloes of all ages, and its immunizing properties were superior to those of the alum-precipitated, agar-wash vaccine and plain bacterin”. An oil adjuvant (paraffin/lanolin) vaccine made from the Katha strain of Z? multocida under field conditions protected 99.6% of vaccinated cattle against HS“‘. Adjuvanted

vis-his

other vaccines

Jayaraman et al.35 vaccinated buffalo calves with alum- precipitated and oil adjuvant vaccine (OAV), and OAV was found to be superior. The superiority of OAV has also been confirmed in the former USSR3h. The efficacies of alum-precipitated broth vaccine and OAV were compared, and it was found that alumprecipitated vaccine offered immunity for 6 months, whereas OAV was found to be a superior vaccine”‘. The OAV does not deteriorate when stored at room temperature for 1 year and can withstand fairly high temperatures. The advantage of its transport without any loss of potency has been responsible for its popularity and success in India”‘.“‘. The vaccine stored at 45°C for about 800 days and at 37-42°C for 20 days still conferred immunity in vaccinated cattle”“. OAV has also been reported to provide immunity for 18 months”,“’ and up to 26 months in cattle”‘. Water buffaloes have also been successfully vaccinated with OAV’j. Immunity was conferred for at least 35 days following vaccinationj4. Two oil adjuvant vaccines of l? multocida Robert type-1 were prepared. The first was a water-in-oil emulsion and contained Marco1 52, Montain de 103 and an antigen ratio of 6:1:3, and the second was a double emulsion containing Marco1 52, Ailcel A and Tween 80 besides antigen. Both preparations induced sustained high antibody titres in buffalo calves beyond 230 days post-vaccination”. Two FAO

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Haemorrhagic septicaemia vaccines: R. Verma and T.N. Jaiswal International Meetings on HS (1962) recommended that killed cultures with adjuvants are the vaccines of choice, and the aerated broth method is the best technique for its production. This was also confirmed by the APHCA Workshop on HS held in Sri Lanka in 1979. The main problems with adjuvanted vaccine seem to be the extent of local tissue and inflammatory response4h.47. Various oil emulsions in vaccines have been used”. Sodium alginate and OAV gave comparable protection, but the former was easier to prepare and easier to administer4”. Comparisons of the complete and incomplete Freund’s adjuvant with oil adjuvants showed that a higher titre was obtained with Freund’s adjuvant”“. Evaluation of various adjuvants, viz. aluminium hydroxide, oil adjuvant, multiple emulsion and sodium alginate in HS vaccines showed that the oil adjuvant and multiple emulsion were good adjuvants for protectier?‘. In Vietnam, an incomplete seppic adjuvant in an oil base, named Montamide ISA-50, incorporating formolin-killed vaccine has been found similar to OAV. This vaccine was stable for 12 months at 4°C and 37°C possessed a low viscosity, and hence did not produce a local swelling in cattle and buffalo”.““. Multiple

emulsion

(ME) vaccine

The OAV possess a thick viscosity, and it is therefore difficult to administer, sometimes causing swelling at the site of inoculatior?. A multiple emulsion (ME) vaccine prepared by re-emulsifying the OAV with 2% Tween SO was effective and was easy to administer”‘.‘“. Mittal et a1.53.s4inoculated 2 ml and 6 ml of the ME vaccine intramuscularly in rabbits and buffalo, respectively. They reported protection in calves up to 9 months assessed by the direct challenge test. In a comparative efficacy of the oil-adjuvant and multiemulsion oil-adjuvant HS vaccines, both of these vaccines were found to be quite safe in a field trial in 1200 dairy cattle and immunogenic to 100 calves when assessed by a passive mouse protection test and in rabbits by a direct challenge test. The duration of immunity afforded by multiemulsion HS vaccine was slightly less than that afforded by a conventional oil-adjuvant vaccine in a limited study carried up to 1 year with samples constituting a ratio of 3:3 and 3:2 (conventional vaccine+Tween SO saline), but was comparable when mixed in the ratio of 3:155. Double emulsion (DE) vaccine has been shown to be effective parallel to OAV and provided immunity in buffalo, lasting up to 12 months post-vaccination. . Maurya” vaccinated buffalo calves intramuscularly with 6 ml of ME vaccine and studied the immunological parameters. Recently, a ME vaccine has been prepared by re-emulsifying the l? multocida (P52) suspension (matching with brown’s opacity tube no. 10) with 2% Tween 80, and cross-bred male calves were immunized intramuscularly with a single dose of 4 ml. Calves were protected for up to 1 year, as assessed by a direct challenge tes?. Low-volume

HS vaccine

A low-volume HS vaccine has been evaluated in rabbit and cattle. Two millilitres of the vaccine given intramuscularly protected the animal, and the vaccine could be administered easily5”.

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PURIFIED VACCINE

CAPSULAR

EXTRACT

Capsular extract has been used to immunize cattle”“, but this work could not result in a practical vaccine. Oil adjuvant-type vaccine using capsular extracts obtained by solvent precipitation from the supernatant fluid of fermenter grown I? multocida type B has been used for immunization but did not yield any encouraging resultsh’.c’2. LIVE VACCINES Attenuated

vaccine

Prolonged in-vitro cultivation of t? multocida strains can lead to considerable attenuation, a fact originally noted by Pasteur”. The first attenuated bacteria as a vaccine was developed” to prevent the disease in fowls, but this had the drawback that the bacteria reverted to their original virulence. Oreste and Armani”” experimented with Pasteurella from barbone and attenuated them by growing at 32°C and passaging them through pigeons. Hudsonh4 isolated blue variants from old aerated cultures and selected strains for their use as live attenuated vaccine for cattle on the basis of their low pathogenicity to mice. Attempts made from time to time to attenuate a F1 multocida culture without affecting its antigenicity by passaging them through rabbits have not been successful.“s Serial passage of the virulent I! multocida (P52) strain of Mukteswar (Uttar Pradesh, India) in a developing chick embryo”” and in albino rats similarly failed to yield any useful results. Chemically

altered live vaccine

Wei and Carter” developed a live streptomycindependent mutant of l? multocida type B that has been shown to be highly immunogenic in mice and rabbits. Similarly, N-methyl-N-nitro-Nnitrosoguandine has been used to modify and improve procedures to obtain streptomycin dependent mutants of l? multocida type A and type 1 l? haemolytica”. Kucera et al.‘” altered I? multocida chemically using different concentrations of diamino acridine salts (acriflavin). This vaccinal strain offered protection in cattle, swine and sheep against challenge. Efficacy to a streptomycin-depenedent serotype of F1 multocida 3:A mutant vaccine in preventing respiratory pasteurellosis in rabbits after intranasal challenge with virulent wild type F1multocida has been reported’“. A vaccine prepared from live streptomycin-dependent p multocida serotype A: 12 gave disappointing results when used in the control of p muftocida infection in rabbits”. A live vaccine of non-replicable p multocida F53 cells has been prepared by psoralin treatment and long-wave UV irradiation”. A number of mutants have been prepared from Sri lanka strains of p multocida type 6:B’.. One of these mutants, when administered as a live culture, immunized 75% cattle and 100 buffaloes better in a single dose of 1OY”’organisms. A booster dose given 3 weeks later was found to enhance the immunity in cattle. Live vaccine

A live vaccine based on an antigenically related deer strain of l? multocida (serotype B:3,4) that had been

Haemorrhagic isolated from a fallow deer in England7” protected calves against a challenge with serotype B:2 9 months after vaccination7s. A lyophilized form of the vaccine protected three of five subcutaneously vaccinated cattle and three of four intradermally vaccinated cattle against challenge with t! multocida B:2 12 months later, whereas the controls succumbed”. In an other field trial, 1415 cattle and 303 buffaloes were vaccinated subcutaneously’“. No disease was seen in the subcutaneously vaccinated animals, but three buffaloes died soon after vaccination. Since the subcutaneous route of vaccination in young buffaloes resulted in some deaths’“.“, the effect of the route of vaccination was evaluated by vaccinating 674 cattle and buffaloes intranasally and 8231 subcutaneously”. The intranasal route of vaccination was not protective against the standard subcutaneous challenge techniques. Therefore, after large-scale field use, the safety of this vaccine has been questionable for primary vaccination of young buffaloes7”. This vaccine strain is rough and serologically different from typical HS strains that carry B:2 antigens. The difference in somatic antigens (type 3 and 4 carried by the vaccine strain and type 2 found in HS cultures) seems to be responsible for the reduction in virulence, whereas the identical capsular type B helps in the development of protective immunity against HS7”. In Myanmar, the work on this live vaccine was initiated by Myint and his associates in 1984. This vaccine is used in cattle and buffaloes over 6 months of age”‘. A local reaction, in the form of a lump, was shown in animals vaccinated either subcutaneously or intradermally. Since these routes showed adverse reactions, therefore, the intranasal route is recommended for instillation of this live vaccine, which offers immunity for more than a year’“.““. Temperature-sensitive mutant The isolation and characterization of p multocida type D temperature sensitive (ts) mutants have been describedx’, and a ts mutant of p multocida type B could be produced and used in a live form to rotect P vaccinated calves against homologous challengex-.

OUTER MEMBERANE

PROTEIN (OMP)

Abdullahi et al.*” studied the outer membrane proteins (OMP) profile of l? multocida type A and the related Taxon 13 strains isolated from bovine pneumonia to determine the heterogenicity of this group and to attempt to define a protective immunogen in a mouse septicacmia model by investigation of the immunogenicity of the OMPs. They concluded that in the mouse model of pasteurellosis, the major OMPs were not protective antigens and therefore may be unlikely candidates for vaccines. However, their conclusions appear to contradict the findings of Lu et LI~.~’who showed that 31.5-kDa OMP (presumbaly outer protein) was protective against homologous challenge in a rabbit model, but in a mouse model, protection was afforded against heterologous challenge only if the challenge strain was shown by probing with a monoclonal antibody, to express the 37.5-kDa antigen. Protection was not provided against strains lacking this antigen. They showed that this OMP can be a protective immunogen in the strains that possessed it, but

septicaemia

vaccines:

R. Verma and T.N. Jaiswal

concluded that other antigens may be more important as vaccine candidates, because only 24% of their strains of p multocida expressed this antigen. Out of ten major polypeptides of OMPs in the extract of l? multocidu serotype B:2, immunoblotting showed that the polypeptides with molecular weights of 44, 37 and 30 kDa were the major immunogens. Buffalo calves vaccinated with the OMP vaccine or a commercial HS vaccine developed the highest mean log,,, ELISA titres 21 days post-vaccination. The results suggested that OMP was protective and could be used in vaccines against HS”. they

COMBINED VACCINES Vaccination against pasteurellosis in cattle and sheep has been undertaken in combination with other bacterial vaccines in order to increase the cost efficacy”. Attempts have been made to develop a combined HS and Black Quarter (BQ) vaccine, which was found to confer dependable levels of immunity in cattle”. Bharsefat and Firouzi et ~1.~~ prepared a combined HS and black leg vaccine, which was used in zones where both diseases existed. Ruiz et al.xy prepared a combined formolizcd aluminium hydroxide vaccine with p multocida and parainfluenza-3 virus isolated from a calf. Among 100 calves, those given the vaccine all survived, whereas 80 of the unvaccinated controls died. A comparative study on live heat-sensitive trivalent vaccine containing IBR, PI3 and adenovirus-3 and killed trivalent (IBR, PI3 and Pasteurella) vaccine was used to control some respiratory viral diseases in veal calves”‘. Gugiu et al.” also advocated the use of combined vaccination with Foot and Mouth disease and p multocida. A polyvalent vaccine containing F1 multocida and F1 haemolytica, given intranasally to new-born 3-week and 6-week-old calves, offered immunity. Simultaneous vaccination of cattle with rinderpest and HS vaccines with other respiratory disease complex antigens has been recommended”,‘“.

EXPERIMENTAL VACCINE

ANTI-IDIOTYPE

An anti-idiotype stretegy was employed that showed that polyclonal anti-idiotype antibodies could be produced that could mimic a linear p multocida lipopolysaccharide molecule. The antibodies, when used as a vaccine antigen, induced antibodies that recognized lipopolysaccharide (LPS) and imparted acquired protection when syngenic vaccinates were challenged with homologous organisms’J.

AUTOGENOUS

VACCINE

Marschang et al.“’ developed a formaldehyde-inactivated autogenous vaccine using unspecified Pasteurella isolated from lung tissue or nasal swab of cattle. They inoculated the vaccine into 1185 cattle; of these, 740 developed respiratory symptoms, otherwise the vaccine yielded good results. Furthermore, an outbreak of severe disease of multiple etiology with respiratory symptoms dominant among cattle of all ages was controlled by using autogenous vaccines prepared

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against Pasteurella species present in nasal 200 cattle were vaccinated in this waygh.

swabs, and

IMMUNE AND PROTECTIVE MECHAMISMS Natural immunity Naturally acquired immunity to HS has been describedx,‘0.97. This immunity to HS occurs in approximately ten of the Asian buffalo and cattle. In Sri Lanka, where high, moderate and low HS incidence areas can be identified, the porportion of naturally acquired immunity comes from sub-clinical infection. There are some reports, for example in Chad, where animals are found with a naturally acquired immunity in the absence of any clinical cases. Occassionally, cattle have been found in Australia with a mouse protective antibody to 6:B and also in USA9*. Antibodies against p multocida capsular types B and E, which cause HS, were demonstrated in a high percentage of sera from domestic feeder calves that have not been vaccinated with any Pasteurella organisms. These antibodies were considered to be naturally acquired”‘. Naturally acquired antibody has been demonstrated in sera from buffalo and Zebu cattle in HS enzootic areas of Asia and Africa”~““‘. The immunity has been attributed due to protective antibodies that develop following non-fatal exposure and can persist for more than 1 year in some animals”“.

Non-specific

immunity

There has been little work done on the surface components of R multocida that contribute to interference with ingestion by phagocytosis. Maheswaran and Theis”” studied phagocytosis of an uncapsulated E! multocida strain with that of an encapsulated type B strain and an unencapsulated type A strain by qualitatively measuring the uptake by bovine neutrophils (3H) thymidine-labelled bacteria in the presence of heat stable and heat-labile opsonins. The uncapsulated strain was completely phagocytosed by normal and heat-inactivated bovine serum, which indicated that heat-labile factors in serum (complement) were not involved in opsonization. Encapsulated capsular type B strain was found to interfere with opsonization with normal serum. Bhatnagar et al.“” reported that congglutinin or heterohaemagglutinin against erythrocytes of hill bull, buffalo and goat could be demonstrated in any sera of buafflo calves vaccinated OAV against HS. While testing the efficacy of alum-precipitated bacterin that provided protection to challenge, complement-fixation titres against whole cell bacteria, but not titres against saline capsular extract, were closely correlated with resistance2h. Cytophilic and opsonin adhering antibodies were produced, and their levels rose in rabbits immunized with HS OAV and challenge infected with l! multocidato4.

Humoral and cell-mediated immune responses The protective immune response to HS vaccination has long been attributed to humoral response and can

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be transferred to native animals with serum from vaccinated aniamls’04~‘05. Outside the passive protective effect, very little is known about the kinetics of antibody response following OAV or bacterin or even what constitutes a protective response. Even less is known about the specificity of the antibody response or the functional classes of antibody involved in protection. The immune response to vaccination in animals has mostly been studied on the basis of data on morbidity/ survival 54~5h~“‘h~‘07. The humoral response in a mouse protection test has been widely used as a relatively reliable assessment of immunity in cattle and buffalo”‘.“&‘“. Vaccination of animals elicits a humoral and the presence of circulating immune response, antibody in cattle and buffalo correlates with immunity”“,lox. Dhanda”” showed that levels of circulating antibodies were detectable from the first week up to 18 months post-vaccination with OAV. It has been difficult to determine whether these protective antibodies operated by a bactericidal or as an opsonizing mechanism”‘“-“‘. ELISA and immunoblotting techniques used to examine the humoral immune response to Pasteurella multocida in bovine sera from Indonesia and Malaysia showed elevated levels of antibody in vaccinated animals. Antibodies from the vaccinated cattle strongly reacted with five or six of the 40 protein bands in this organism”‘. An increase in the IgM and IgG, even though the differences were not statistically significant, was reported in calves immunized with a live vaccine containing F1 multocida A:3 and l? haemolytica A”‘. Serum from some cattle vaccinated with the Katha strain or local strains of F1 multocida in oil adjuvant had elevated ELISA values to heat stable LPS antigen”“. The serum antibody response of buffaloes immunized with three conventional HS vaccines, viz. broth bacterin (BB), alum-precipitated vaccine (APV), and oil adjuvant vaccine (OAV) and one experimental double emulsion vaccine (DEV), was determined by ELISA. Antibody levels were significantly higher in buffaloes immunized with the adjuvanted vaccines (APV, OAV, DEV) than those immunized with BB alone. There also appeared to be a relationship between ELISA titres and active protection in buffaloes. A preliminary analysis of the antibody isotype in of buffaloes immunized with the the sera oil-adjuvanted vaccines revealed the antipasteurella activity to be associated predominantly with IgGl or IgG2 isotypes. The principle response to OAV and bacterin vaccination was the 1gG response with only a mild and transient IgM response, and the major protective role was due to the IgG antibody clas?‘. There appeared to be an association bctwecn pre-challenge antibody titres and protection in buffaloes following challenges”. Buffaloes with high antibody levels were protected, whereas those buffaloes with low levels succumbed to challenge. A study on the use of an ELISA serotype B in vaccinated cattle and buffaloes showed that ELISA results were correlated with direct No minimum antibody level necessary for challenge”‘. protection has been attributed because of the difference in the types of methods of antigen preparation”4. Verma and Jaiswal” measured the antibody response by indirect haemagglutination assay (IHA) and ELISA

Haemorrhagic septicaemia vaccines: R. Verma and T.N. Jaiswal in cross-bred male calves immunized with multiple emulsion (ME) HS vaccine. In their study, an animal with a minimum of 80lHA prechallenge titre withstood the challenge infection with virulent P multocida. However, because of the variation in the prechallenge titres among the group of animals, no minimum level of prechallenge antibody titre could be proposed. There is no evidence of a major role of a cellmediated (CMI) immune response in HS”“. However, the involvement of cell- mediated immunity cannot be discounted in protection”“. One of the reasons for not studying the role of cell-mediated immune response in cattle is that P nzultocida has not been described as a facultative intracellular bacteria. The other reason for discounting the cell-mediated response is largely a humoral response in HS. Maheswaran and Theis”” reported that lymphocytes from cattle immunized with various HS strains of l? multocida showed higher stimulation indices when incubated with homologous antigen, which suggested an involvement of cellmediated immunity. The leucocyte migration-inhibition (LMIT) test has been described as an indicator of the cell-mediated immune response”‘. Verma and Jaiswal” reported a percentage migration-inhibition of more than 20 in calves immunized with ME vaccine both at prechallenge and post-challenge intervals, thus indicating the involvement of cell-mediated immune mechanisms. An apparent relationship between delayed type hypersensitivity (DTH) response and protection in buffalo vaccinated with different HS vaccines has been reported.”

VACCINATION

PROGRAMMES

The vaccination programme against HS varies. Vaccination before the onset of a monsoon or sometimes at the onset of the outbreak of the disease is practised in most countries. Table 1 gives information on the type of

Table 1

Durationof

vaccines, their dose, route and duration of immmunity used in different countries. Critical appraisal

Since the discovery of the attenuation of R muftocida responsible for fowl cholera and the first use of bacterin to prevent HS, more than a decade has passed in which numerous efforts have been made to develop a suitable and cost-effective HS vaccine using whole inactivated sells or capsular extract with or without adjuvants. The ultimate success in HS vaccinology has, so far, been the development of oil adjuvant vaccinf?‘,“, and now recently, a live vaccine is used in Myanmar79.x”.Both of these vaccines offer immunity for 1 year, but large-scale vaccination of animals with live HS vaccination in enzootic areas in Myanmar provided immunity over 2years to natural infection in vaccinated animals. Myin? contended that the respiratory tract is considered the natural route of entry of Pasteurells. The organisms that cause HS are carried by healthy animals in their nasopharynx, and such animals are considered as seeds of HS outbreak. As such, local secretory antibodies (specific IgA) are produced due to intranasal vaccination with live HS vaccine, which may be an advantage in preventing natural infection of HS. It appears that slightly more work is needed to elucidiate whether local secretory antibodies are protective. Pa.steurella causes systemic bacteremia and the extent to which this intranasal vaccination will provide protection over the protection provided by circulating IgG needs to be studied in detail. An other success in vaccinology has been the development of a double emulsion vaccine and a multiple emulsion vaccine, which provided immunity for 52 weeks and 1 year, respectively, but these experimental vaccines are yet to go for field use. It has been established in epidemiological studies of P multocida that in-viva antigens are important

immunity offered by HS vaccine used in different countries

Country

Type of vaccine

Dose

Route

Immunity

India

3 3 3 3

ml ml ml ml

SIC SIC

Indonesia

Alum precipitated vaccine Alumninium hydroxide gel vaccine Oil adjuvant vaccine Oil adjuvant vaccine

Thailand

Aluminium

gel

3 ml

SIC

Malaysia

Alum-precipitated

vaccine

5 ml

SIC

Vietnam

Oil adjuvant vaccine Alum-precipitated vaccine

3 ml 2 ml

IIM SIC

2-3 ml 2 ml 3 ml

s/c

Sri Lanka

Aluminium hydroxide Oil adjuvant vaccine Oil adjuvant vaccine

Cambodia Myanmar Phillipines Nepal Iran

Alum adjuvant vaccine Live vaccine Killed alum adjuvanted vaccine Alum-precipitated vaccine HS and Black leg combined vaccine

2 ml 2 ml 5 ml

SIC I/N

4-6 months 4-6 months 1 year or more Animal older than 5 months vaccinated every 6 months Immunity 1 year Vaccine contains 3 x 10” c.f.u. per dose of P. multocida serotype B:2 Annual revaccination Immunity 4-6 months Immunity 1 year Two injections per year Immunity 4-6 months Immunity 4-6 months Immunity 1 year Cattle and buffalo vaccinated at 4-6 months age and given a booster 3-6 months later followed by annual vaccination 3-6 months later twice yearly One dose of serotype B;3,4 Booster every 6 months 6 months

hydroxide

absorbed

vaccine

I/M I/M

I/M I/M

SIC SIC

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Reference

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Haemorrhagic septicaemia vaccines: R. Verma and T.N. Jaiswal immunogens and are responsible for a high level of immunity following arrested infection’27”2x. This observation warrants a need to develop a safe, avirulent, stable variant strain that should produce the important immunogens in vivo when administered as a live vaccine. This would require identification of the important immunogens in viva when administered as a live vaccine. This would require identification of the in-vivo important immunogens responsible for the high level of immunity attained in natural exposure and development of artificial media, which should support in-vitro expression of these immunogens. Adjuvants have played a great role in the development of HS vaccine, and the duration of immunity depends on the type of adjuvant used. The use of adjuvants, including the new generation adjuvants, should be evaluated to give an easily injectible vaccine. Although no recombinant or subunit vaccine against HS is available, in the modern sense of vaccination, future research may be directed to develop such vaccine(s). Immune mechanisms in HS have been poorly understood. Investigations into the immune mechanisms, virulence determinants, comparison of live vs. killed vaccine and the carrier state are some important areas that have been suggested as future lines of research12’.

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