The immune response to bovine viral diarrhea virus: A constantly changing picture

Veterinary Clinics of North America - Food Animal Practice

Volumn 20, Issue 1, 2004, Pages 95-114

  Chase, Christopher C L ^a   Elmowalid, Gamal ^a,b   Yousif, Ausama A A ^a,c  

^a SOUTH DAKOTA STATE UNIVERSITY   (United States)

^b Zagazig University   (Egypt)

^c Cairo University   (Egypt)

Author keywords

[No Author keywords available]

Indexed keywords

ALPHA INTERFERON; DNA VACCINE; GAMMA INTERFERON; INACTIVATED VIRUS VACCINE; LYMPHOTOXIN; NEUTRALIZING ANTIBODY; TUMOR NECROSIS FACTOR ALPHA; VIRUS VACCINE;

ANTIGEN PRESENTATION; BOVINE DIARRHEA VIRUS; CATTLE DISEASE; CELLULAR IMMUNITY; CROSS REACTION; CYTOKINE PRODUCTION; CYTOTOXIC T LYMPHOCYTE; DRUG EFFECT; GRANULOCYTE; HELPER CELL; HUMORAL IMMUNITY; IMMUNE RESPONSE; INFECTION PREVENTION; MACROPHAGE; MONOCYTE; NATURAL KILLER CELL; NONHUMAN; PERSISTENT INFECTION; PHAGOCYTOSIS; REVIEW; T LYMPHOCYTE SUBPOPULATION; VACCINATION; VIRUS INFECTION; VIRUS REPLICATION; VIRUS STRAIN;

ANIMALS; ANTIBODIES, VIRAL; ANTIGENS, VIRAL; BOVINE VIRUS DIARRHEA-MUCOSAL DISEASE; CATTLE; DIARRHEA VIRUSES, BOVINE VIRAL; IMMUNITY, CELLULAR; IMMUNITY, MATERNALLY-ACQUIRED; IMMUNITY, NATURAL; MACROPHAGES; VIRAL VACCINES;

BOS TAURUS; BOVINAE; BOVINE VIRAL DIARRHEA VIRUS 1;

EID: 2342501369     PISSN: 07490720     EISSN: None     Source Type: Journal    
DOI: 10.1016/j.cvfa.2003.11.004     Document Type: Review

References (101)

1. Potgieter LN. Immunology of bovine viral diarrhea virus. Vet Clin North Am Food Anim Pract 1995;11(3):501-20.
2. Glew EJ, Carr BV, Brackenbury LS, Hope JC, Charleston B, Howard CJ. Differential effects of bovine viral diarrhoea virus on monocytes and dendritic cells. J Gen Virol 2003;84(7):1771-80.
3. Lambot M, Hanon E, Lecomte C, Hamers C, Letesson JJ, Pastoret PP. Bovine viral diarrhoea virus induces apoptosis in blood mononuclear cells by a mechanism largely dependent on monocytes. J Gen Virol 1998;79(5):1745-9.
4. Archambault D, Beliveau C, Couture Y, Carman S. Clinical response and immunomodulation following experimental challenge of calves with type 2 noncytopathogenic bovine viral diarrhea virus. Vet Res 2000;31:215-27.
5. Peterhans E, Jungi TW, Schweizer M. BVD virus and innate immunity. Biologicals 2003;31:107-12.
6. Welsh MD, Adair BM, Foster JC. Effect of BVD virus infection on alveolar macrophage functions. Vet Immunol Immunopathol 1995;46:195-210.
7. Liu L, Lemkuhl HD, Kaeberle ML. Synergistic effects of bovine respiratory syncytial virus and non-cytopathic bovine viral diarrhea virus infection on selected bovine alveolar macrophage functions. Can J Vet Res 1999;63:41-8.
8. Olchowy TWJ, Slauson DO, Boschsler PN. Induction of procoagulant activity in virus infected bovine alveolar macrophages and the effect of lipopolysaccharide. Vet Immunol Immunopathol 1997;58:27-37.
9. Adler H, Frech B, Meier P, Jungi TW, Peterhans E. Noncytopathic strains of bovine viral diarrhea virus prime bovine bone marrow-derived macrophages. Biochem Biophy Res Commun 1994;202:1562-8.
10. Adler H, Jungi TW, P.ster H, Strasser M, Sileghem M, Peterhans E. Cytokine regulation by virus infection: bovine viral diarrhea virus, a flavivirus, downregulates production of tumor necrosis factor alpha in macrophages in vitro. J Virol 1996;70:2650-3.
11. Adler B, Adler H, Pfeister H, Jungi TW, Peterhans E. Macrophages infected with cytopathic bovine viral diarrhea virus release a factor(s) capable of priming uninfected macrophages for activation-induced apoptosis. J Virol 1997;71:3255-8.
12. Van Reeth K, Adair B. Macrophages and respiratory viruses. Pathol Biol 1997;45:184-92.
13. rd congress of the European Society of Veterinary Virology, Interlaken; 1995. p. 63-7.
14. Jensen J, Schultz RD. Effect of infection by bovine viral diarrhea virus (BVDV) in vitro on interleukin-1 activity of bovine monocytes. Vet Immunol Immunopathol 1991;29:251-65.
15. Ketelsen AT, Johnson DW, Muscoplat CC. Depression of bovine monocyte chemotactic responses by bovine viral diarrhoae virus. Infect Immun 1979;25:225-30.
16. Teichmann U, Liebler-Tenorio E, Pohlenz JF. 2000. Ultrastructural changes in follicles of small-intestinal aggregated lymphoid nodules in early and advanced phases of experimentally induced mucosal disease in calves. Am J Vet Res 2000;61:174-82.
17. Liebler-Tenorio EM, Ridpath JF, Neill JD. Lesions and tissue distribution of viral antigen in severe acute versus subclinical acute infection with BVDV2. Biologicals 2003;31:119-22.
18. Liebler-Tenorio EM, Ridpath JF. Distribution of viral antigen and development of lesions after experimental infection with highly virulent bovine viral diarrhea virus type 2 in calves. Am J Vet Res 2002;63:1575-84.
19. Stoffregen B, Bolin SR, Ridpath JF, Pohlenz J. Morphologic lesions in type 2 BVDV infections experimentally induced by strain BVDV2-1373 recovered from a field case. Vet Microbiol 2000;77:157-62.
20. Sentsui H, Takami R, Nishimori T, Murakami K, Yokoyama T, Yokomizo Y. Anti-viral effect of interferon-α on bovine viral diarrhea virus. J Vet Med Sci 1998;60:1329-33.
21. Perler L, Schweizer M, Jungi TW, Peterhans E. Bovine viral diarrhea virus and bovine herpesvirus-1 prime uninfected macrophages for lipopolysaccharide- triggered apoptosis by interferon-dependent and -independent pathways. J Gen Virol 2000;81:881-7.
22. Schweizer M, Peterhans E. Noncytopathic bovine viral diarrhea virus inhibits doublestranded RNA-induced apoptosis and interferon synthesis. J Virol 2001;75:4692-8.
23. Charleston B, Fray MD, Baigent S, Carr BW, Morrison WI. Establishment of persistent infection with non-cytopathic bovine viral diarrhea virus in cattle is associated with a failure to induce type I interferon. J Gen Virol 2001;82:1893-7.
24. Charleston B, Brackenbury LS, Carr BV, Fray MD, Hope JC, Howard CJ, et al. Alpha/beta and gamma interferons are induced by infection with noncytopathic viral diarrhea virus in vivo. J Virol 2002;76:923-7.
25. Brodersen BW, Kelling CL. Alteration of leukocyte populations in calves concurrently infected with bovine respiratory syncytial virus and bovine viral diarrhea virus. Viral Immun 1999;12:323-34.
26. Ellis JA, Davis WC, Belden EL, Pratt DL. Flow cytofluorimetric analysis of lymphocyte subset alterations in cattle infected with bovine viral diarrhea virus. Vet Pathol 1988;25:231-6.
27. Howard CJ, Clarke MC, Sopp P, Brownlie J. Immunity to bovine virus diarrhoea virus in calves: the role of different T-cell subpopulations analysed by specific depletion in vivo with monoclonal antibodies. Vet Immunol Immunopathol 1992;32:303-14.
28. Bruschke CJM, Haghparast A, Hoek A, Rutten VPMG, Wentink GH, van Rijn PA, et al. The immune response of cattle, persistently infected with noncytopathic BVDV, after superinfection with antigenically semi-homologous cytopathic BVDV. Vet Immunol Immunopathol 1998;62:37-50.
29. Lambot M, Douart A, Joris E, Letesson JJ, Pastoret PP. Characterization of the immune response of cattle against non-cytopathic and cytopathic biotypes of bovine viral diarrhoea virus. J Gen Virol 1997;78:1041-7.
30. Collen T, Morrison WI. CD = 4 T-cell responses to bovine viral diarrhoea virus in cattle. Virus Res 2000;67:67-80.
31. + T-cell recognition of a heterologous virus by cattle persistently infected with bovine viral diarrhea virus. Virology 2000;276:70-82.
32. + 4 T cells reactive with bovine diarrhoea virus. Vet Immunol Immunopathol 2002;87:235-8.
33. Waldvogel AS, Hediger-Withaler BM, Eicher R, Zakher A, Zarlenga DS, Gasbarre LC, et al. Interferon-γ and interleukin-4 mRNA expression by peripheral blood mononuclear cells from pregnant and non-pregnant cattle seropositive for bovine viral diarrhea virus. Vet Immunol Immunopathol 2000;77:201-12.
34. Rhodes SG, Cocksedge JM, Collins RA, Morrison WI. Differential cytokine responses of CD4+ and CD8 + T cells in response to bovine viral diarrhea virus in cattle. J Gen Virol 1999;80:1673-9.
35. Hou I, Wilkerson M, Kapil S, Mosier D, Shuman W, Reddy JR, et al. The effect of different viral diarrhea virus genotypes and biotypes on the metabolic activity and activation status of bovine peripheral blood mononuclear cells. Viral Immunol 1998;11:233-44.
36. Charleson B, Hope JC, Carr BV, Howard CJ. Masking of two in vitro immunological assays for Mycobacterium bovis (BCG) in calves acutely infected with non-cytopathic bovine viral diarrhoea virus. Vet Rec 2001;149:481-4.
37. Beer M, Wolf G, Pichler A, Wolfmeyer A, Kaaden OR. Cytotoxic T-lymphocyte responses in cattle infected with bovine viral diarrhea virus. Vet Micro 1997;58:9-22.
38. Hegde N, Srikumaran S. The use of bovine MHC class I allele-specific peptide motifs and proteolytic cleavage specificities for the prediction of potential cytotoxic T lymphocyte epitopes of bovine viral diarrhea virus. Virus Genes 1997;14:111-21.
39. Donis RO. Molecular biology of BVDV and its interaction with the host. Vet Clin North Am Food Anim Pract 1995;11:393-423.
40. Deregt D, van Rijn PA, Wiens TY, van den Hurk J. Monoclonal antibodies to the E2 protein of a new genotype (type 2) of bovine viral diarrhea virus define three antigenic domains involved in neutralization. Virus Res 1998;57:171-81.
41. Deregt D, Bolin SR, van den Hurk J, Ridpath JF, Gilbert SA. Mapping of a type 1-specific and a type-common epitope on the E2 (gp53) protein of bovine viral diarrhea virus with neutralization escape mutants. Virus Res 1998;53:81-90.
42. Vilcek S. Identification of pestiviruses contaminating cell lines and fetal calf sera. Acta Virol 2001;45:81-6.
43. Nobiron I, Thompson I, Brownlie J, Collins ME. DNA vaccination against bovine viral diarrhoea virus induces humoral and cellular responses in cattle with evidence for protection against viral challenge. Vaccine 2003;21:2082-92.
44. Lambot M, Joris E, Douart A, Lyaku J, Letesson JJ, Pastoret PP. Evidence for biotype-specific effects of bovine viral diarrhoea virus on biological responses in acutely infected calves. J Gen Virol 1998;79:27-30.
45. Chase CC, Chase SK, Fawcett L. Trends in the BVDV serological response in the upper midwest. Biologicals 2003;31:145-51.
46. Fredriksen B, Sandvik T, Loken T, Odegaard SA. Level and duration of serum antibodies in cattle infected experimentally and naturally with bovine virus diarrhoea virus. Vet Rec 1999;144:111-4.
47. Hamers C, di Valentin E, Lecomte C, Lambot M, Joris E, Genicot B, et al. Virus neutralizing antibodies against a panel of 18 BVDV isolates in calves vaccinated with Rispoval RS-BVD. J Vet Med B Infect Dis Vet Public Health 2000;47:721-6.
48. Cortese VS, Whittaker R, Ellis J, Ridpath JF, Bolin SR. Specificity and duration of neutralizing antibodies induced in healthy cattle after administration of a modified-live virus vaccine against bovine viral diarrhea. Am J Vet Res 1998;59:848-50.
49. Hamers C, di Valentin E, Lecomte C, Lambot M,Joris E, Genicot B, et al. Virus neutralizing antibodies against a 22 bovine isolates in vaccinated calves. Vet J 2002;163:61-7.
50. Jones L, Van Campen H, Zu ZC, Schnackel JA. Comparison of neutralizing antibodies to type 1a, 1b and 2 bovine viral diarrhea virus from experimentally infected and vaccinated cattle. Bovine Practitioner 2001;35:137-40.
51. Fulton RW, Ridpath JF, Saliki JT, Briggs RE, Confer AW, Burge LJ, et al. Bovine viral diarrhea virus (BVDV) 1b: predominant BVDV subtype in calves with respiratory disease. Can J Vet Res 2002;66:181-90.
52. Fulton RW, Ridpath JF, Confer AW, Saliki JT, Burge LJ, Payton ME. Bovine viral diarrhoea virus antigenic diversity: impact on disease and vaccination programmes. Biologicals 2003;31:89-95.
53. Fulton RW, Burge LJ. Bovine viral diarrhea virus types 1 and 2 antibody response in calves receiving modified live virus or inactivated vaccines. Vaccine 2001;19:264-74.
54. Cortese VS, West KH, Hassard LE, Carman S, Ellis JA. Clinical and immunologic responses of vaccinated and unvaccinated calves to infection with a virulent type-II isolate of bovine viral diarrhea virus. J Am Vet Med Assoc 1998;213:1312-9.
55. Ellis J, West K, Cortese V, Konoby C, Weigel D. Effect of maternal antibodies on induction and persistence of vaccine-induced immune responses against bovine viral diarrhea virus type II in young calves. J Am Vet Med Assoc 2001;219:351-6.
56. Munoz-Zanzi CA, Thurmond MC, Johnson WO, Hietala SK. Predicted ages of dairy calves when colostrum-derived bovine viral diarrhea virus antibodies would no longer offer protection against disease or interfere with vaccination. J Am Vet Med Assoc 2002;221:678-85.
57. Ridpath JF, Neill JD, Endsley J, Roth JA. Effect of passive immunity on the development of a protective immune response against bovine viral diarrhea virus in calves. Am J Vet Res 2003;64:65-9.
58. Endsley J, Roth JA, Ridpath J, Neill J. Maternal antibody blocks humoral but not T cell responses to BVDV. Biologicals 2003;31:123-5.
59. Brownlie J, Hooper LB, Thompson I, Collins ME. Maternal recognition of foetal infection with bovine virus diarrhoea virus (BVDV)-the bovine Pestivirus. Clin Diag Virol 1998;10:141-50.
60. Swasdipan S, McGowan M, Phillips N, Bielefeldt-Ohmann H. Pathogenesis of transplacental virus infection: pestivirus replication in the placenta and fetus following respiratory infection. Micro Pathol 2002;32:49-60.
61. Glew EJ, Howard CJ. Antigen-presenting cells from calves persistently infected with bovine diarrhoea virus, a member of the Flaviviridae, are not compromised in their ability to present viral antigen. J Gen Virol 2001;82:1677-85.
62. Collins ME, Desport M, Brownlie J. Bovine viral diarrhea quasispecies during persistent infection. Virology 1999;259:85-98.
63. Fray MD, Supple EA, Morrison WI, Charleston B. Germinal center localization of bovine viral diarrhoea virus in persistently infected animals. J Gen Virol 2000;81:1669-73.
64. Fulton RW, Step DL, Ridpath JF, Saliki JT, Confer AW, Johnson BJ, et al. Response of calves persistently infected with noncytopathic bovine viral diarrhea virus (BVDV) subtype 1b after vaccination with heterologous BVDV strains in modified live virus vaccines and Mannheimia haemolytica bacterin-toxoid. Vaccine 2003;21:2980-5.
65. van Oirschot JT, Bruschke CJM, van Rijn PA. Vaccination of cattle against bovine viral diarrhoea. Vet Microbiol 1999;64:169-83.
66. AgriLaboratories. Immunogenicity of titanium 5 L5 following BVDV 890 (type 2) challenge. Challenge study test results, TR-01. St. Joseph (MO): AgriLabs; 1998.
67. AgriLaboratories. Immunogenicity of titanium 5 L5 following BVDV NY-1 (type 1) challenge. Challenge study test results, TR-02. St. Joseph (MO): AgriLabs; 1999.
68. AgriLaboratories. Immunogenicity of master guard following type 2 challenge. Challenge study test results, TR-06. St. Joseph (MO): AgriLabs; 1999.
69. Boehringer Ingelheim Vetmedica. BVD vaccination and challenge trials for BVD types 1 and 2 vaccines. Technical bulletin TB99-107. St. Joseph (MO): Boehringer Ingelheim Vetmedica; 1999.
70. Endsley JJ, Quade MJ, Terhaar B, Roth JA. Bovine viral diarrhea virus type 1- and type 2-specific bovine T lymphocyte-subset responses following modified-live virus vaccination. Vet Ther 2002;3:364-72.
71. Fairbanks K, Schnackel J, Chase CC. Evaluation of a modified live virus type-1a bovine viral diarrhea virus vaccine (Singer Strain) against a type-2 (strain 890) challenge. Vet Ther 2003;4:24-34.
72. Dean HJ, Leyh R. Cross-protective efficacy of a bovine viral diarrhea virus (BVDV) type 1 vaccine against BVDV type 2 challenge. Vaccine 1999;17:1117-24.
73. Makoschey B, Janssen MGJ, Vrijenhoek MP, Korsten JHM, Marel PVD. An inactivated bovine virus diarrhoea virus (BVDV) type 1 vaccine affords clinical protection against BVDV type 2. Vaccine 2001;19:3261-8.
74. Van Campen H, Vorpahl P, Huzurbazar S, Edwards J, Cavender J. A case report: evidence for type 2 bovine viral diarrhea virus (BVDV)-associated disease in beef herds vaccinated with a modified-live type 1 BVDV vaccine. J Vet Diag Invest 2000;12:263-5.
75. Thurmond MC, Muñoz-Zanzi CA, Hietala SK. Effect of calfhood vaccination on transmission of bovine viral diarrhea virus under typical drylot dairy conditions. JAVMA 2001;219:968-75.
76. Brownlie J, Clarke MC, Hooper LB, Bell GD. Protection of the bovine fetus from bovine viral diarrhoea virus by means of a new inactivated vaccine. Vet Rec 1995;137:58-62.
77. Dean HJ, Hunsaker BD, Bailey OD, Wasmoen T. Prevention of persistent infection in calves by vaccination of dams with noncytopathic type-1 modified-live bovine viral diarrhea virus prior to breeding. Am J Vet Res 2003;64:530-7.
78. Cortese VS, Grooms DL, Ellis J, Bolin SR, Ridpath JF, Brock KV. Protection of pregnant cattle and their fetuses against infection with bovine viral diarrhea virus type-1 by use of a modified-live virus vaccine. Am J Vet Res 1998;59:1409-13.
79. Brock KV, Cortese VS. Experimental fetal challenge using type II bovine viral diarrhea virus in cattle vaccinated with modified-live virus vaccine. Vet Ther 2001;2:354-60.
80. Paton DJ, Sharp G, Ibata G. Foetal cross-protection experiments between type 1 and type 2 bovine viral diarrhoea virus in pregnant ewes. Vet Microbiol 1999;61:185-96.
81. Patel JR, Shilleto RW, Williams J, Alexander DCS. Prevention of transplacental infection of bovine foetus by bovine viral diarrhoea virus through vaccination. Arch Virol 2002;147:2453-63.
82. Bruschke CJM, van Oirschot JT, van Rijn PA. An experimental multivalent bovine virus diarrhea virus E2 subunit vaccine and two experimental conventionally inactivated vaccines induce partial fetal protection in sheep. Vaccine 1999;17:1983-91.
83. Zimmer GM, Wentink GH, Bruschke C, Westenbrink FJ, Brinkhof J, de Goey I. Failure of foetal protection after vaccination against an experimental infection with bovine viral diarrhea virus. Vet Micro 2002;89:255-65.
84. Harpin S, Talbot B, Mbikay M, Elazhary Y. Immune response to vaccination with DNA encoding the bovine viral diarrhea virus major glycoprotein gp53 (E2). FEMS Microbiol Lett 1997;146:229-34.
85. Harpin S, Hurley DJ, Mbikay M, Talbot B, Elazhary Y. Vaccination of cattle with a DNA plasmid encoding the bovine viral diarrhea virus major glycoprotein E2. J Gen Virol 1999;80:3137-44.
86. Nobiron I, Thompson I, Brownlie J, Collins ME. Co-adminstration of IL-2 enhances antigen-specific immune responses following vaccination with DNA encoding the glycoprotein E2 of bovine viral diarrhoea virus. Vet Micro 2000;76:129-42.
87. Nobiron I, Thompson I, Brownlie J, Collins ME. Cytokine adjuvancy of BVDV DNA vaccine enhances both humoral and cellular immune responses in mice. Vaccine 2001;19:4226-35.
88. Bolin SR, Ridpath JF. Glycoprotein E2 of bovine viral diarrhea virus expressed in insect cells provides calves limited protection from systemic infection and disease. Arch Virol 1996;141:1463-77.
89. Elahi SM, Shen SH, Talbot BG, Massie B, Harpin S, Elazhary Y. Recombinant adenoviruses expressing the E2 protein of bovine viral diarrhea virus induce humoral and cellular immune responses. FEMS Microbiol Lett 1999;177:159-66.
90. Elahi SM, Shen SH, Harpin S, Talbot BG, Elazhary Y. Investigation of the immunological properties of the bovine viral diarrhea virus protein NS3 expressed by an adenovirus vector in mice. Arch Virol 1999;144:1057-70.
91. Elahi SM, Shen SH, Talbot BG, Massie B, Harpin S, Elazhary Y. Induction of humoral and cellular immune responses against the nucleocapsid of bovine viral diarrhea virus by an adenovirus vector with an inducible promoter. Virology 1999;261:1-7.
92. Baxi MK, Deregt D, Robertson J, Babiuk LA, Schlapp T, Tikoo SK. Recombinant bovine adenovirus type 3 expressing bovine viral diarrhea virus glycoprotein E2 induces an immune response in cotton rats. Virology 2000;278:234-43.
93. Toth RL, Nettleton PF, McCrae MA. Expression of the E2 envelope glycoprotein of bovine viral diarrhea virus (BVDV) elicits virus-type specific antibodies. Vet Micro 1999;65:87-101.
94. Bogdan C. Nitric oxide and the immune response. Nat Immunol 2001;2:907-16.
95. Spellberg B, Edwards JE. Type 1/Type 2 immunity in infectious diseases. Clin Infect Dis 2001;32:76-102.
96. Bolin SR. The pathogenesis of mucosal disease. Vet Clin North Am Food Anim Pract 1995;11:489-500.
97. Ridpath JR. BVDV genotypes and biotypes: practical implications for diagnosis and control. Biologicals 2003;31:127-31.
98. National Animal Health Monitoring System (NAHMS). Part III: Reference of 1997 Beef Cow-calf Production Management & Disease Control, 1998; NAHMS, Animal and Plant Health Inspection Service, United States Department of Agriculture. http://www.aphis.usda.gov/vs/ceah/cahm/Beef_Cow-Calf/bf97des3.pdf.
99. National Animal Health Monitoring System (NAHMS). Part 1: Reference of 1996 Dairy Management Practices, 1996; NAHMS, Animal and Plant Health Inspection Service, United States Department of Agriculture. http://www.aphis.usda.gov/vs/ ceah/cahm/ Dairy_Cattle/dr96des1.pdf.
100. National Animal Health Monitoring System (NAHMS). Vaccination Practices for Respiratory Pathogens in US Feedlots, 1995; NAHMS, Animal and Plant Health Inspection Service, United States Department of Agriculture, http://www.aphis.usda.gov/vs/ceah/cahm/Beef_Feedlot/cofvac.pdf.
101. National Animal Health Monitoring System (NAHMS). Vaccination of Cattle Against Respiratory Disease Pathogens in US Feedlots, 2000; NAHMS, Animal and Plant Health Inspection Service, United States Department of Agriculture. http://www.aphis.usda.gov/vs/ceah/cahm/Beef_Feedlot/FD99vacc.htm.