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Introduction
Respiratory Disease
Abortion
Neurological Disease
Disease in Newborn Foals
Prevention & Control
Bibliography

Diseases of the Horse Caused by Equine Herpesvirus Type 1 (EHV-1) - A Bibliography of Source Materials


I. Brief Overview

A. Introduction

Of all the infectious viral diseases recognized in horses, those caused by equine herpesvirus type 1 (EHV-1) (Fig 1) are among the most costly to the equine industry. EHV-1 is unique in its ability to target and attack three separate organ systems of the horse (respiratory, reproductive, and central nervous systems), giving rise to large-scale outbreaks with any permutation of respiratory tract disease, abortion, neonatal mortality, and a paralytic neurological disease. All four EHV-1 disease manifestations continue to plague the equine breeding industry with insidious and cumulatively immense economic and animal welfare losses despite many years of research and the widespread use of numerous vaccines. Clearly, there is much about these herpesvirus diseases yet to be learned before they can be brought under an acceptable level of control, and continued research on EHV-1 has, therefore, been an international equine health priority for many years.

EHV-1 is deeply entrenched within the world’s horse population as inapparent, latent infections (silent carriers). The capacity of EHV-1 to persist in the body of the horse in a dormant, but potentially reactivatable, state after recovery from a primary infection provides an inexhaustible and globally distributed biological reservoir of the virus for continuous transmission of infection among adult horses and to new generations of young horses (Fig 2).

Figure 1. Electron photomicrograph of equine herpesvirus type 1.

photo of ehv1

Figure 2. Epidemiological cycle of EHV-1 infections, illustrating the central role of latently infected carrier horses as biological reservoirs from which the virus is perpetually transmitted to new generations of equine hosts.

 

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B. EHV-1 Respiratory Disease

EHV-1 is among the large spectrum of respiratory pathogens capable of causing clinically significant upper respiratory tract disease in the domestic horse. EHV-1 respiratory disease is associated with destructive infection of the upper airway epithelium and is a condition seen primarily in young horses. A febrile episode accompanied by bilateral nasal discharge, inappetence and depression is the common clinical presentation (Fig 3). While the majority of such respiratory infections by EHV-1 are subclinical or mild in nature, the herpesvirus has the potential for causing widespread outbreaks of severe upper respiratory tract disease. The greatest burden of herpesvirus respiratory disease is borne by young horses, with the window of highest risk between weaning and 3 years of age. In two- and three-year-old horses in training or on the racing circuit, in which horses from diverse origins are housed together in enclosed, confined spaces such as race or show barns, outbreaks of viral respiratory disease are common. Epidemics of EHV-1 upper respiratory tract disease have been described in such settings with some outbreaks resulting in neurological sequelae. Quantitatively, herpesvirus outbreaks generally take second place to equine influenza as a significant threat for acute respiratory disease in two- and three-year-olds. Previously exposed horses older than 3 years of age continue to show serological evidence of periodic re-infection by EHV-1 throughout their lifetimes with only minimal and transient clinical signs of respiratory tract infection. In such animals, however, the risks for the more serious sequelae of abortion and/or neurological disease following subclinical respiratory infection by EHV-1 continue to exist.

 

Figure 3. Nasal discharge in a weanling foal with febrile EHV-1 respiratory tract infection.

photo of respiratory disease

 

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C. EHV-1 Abortion

Abortion in broodmares caused by infection of the fetus with EHV-1 is among the most widespread and economically important, infectious disease problems of the horse (Fig 4). Partial success at controlling the economic losses attributable to EHV-1 has been achieved, after many years of effort, by programs of intensive annual vaccinations. Mares infected with EHV-1 abort precipitously with no impending signs, and evidence of the previous respiratory tract infection is usually not observed. The placenta is expelled along with the fetus that is often still enveloped in its amniotic membrane. At the time of abortion, the fetus has usually just died from asphyxia associated with sudden separation of the placenta from the endometrium that precedes fetal expulsion. The aborted fetus possesses high levels of virus and extensive histopathological evidence of multi-organ infection. Almost all EHV-1 abortions occur during the last four months of gestation. Most abortion occurrences associated with EHV-1 infection involve only one or two mares in a group. However, epidemic abortigenic disease (abortion storms) claiming high percentages of the potential foal crop also occurs. Once a mare has aborted, her future reproductive potential is not compromised; most mares conceive successfully shortly after abortion and foal normally the following year.
Figure 4. Equine fetus aborted as a result of infection by equine herpesvirus-1. The fetus is still attached to the placenta and enclosed in its amnion.

photo of aborted equine fetus

 

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D. EHV-1 Neurological Disease

The clinical outcome of EHV-1 infection that carries the most dramatic negative consequences for the welfare of horses is the dreaded neurological disease. Once regarded as a rare and sporadic occurrence, the malady has in recent years been observed with increasing and alarming frequency. Common to all horses with EHV-1 associated central nervous system disease are a weakness and paralysis of the muscles of the hind limbs that give rise to incoordination, gait abnormalities and, in many cases, the inability to rise from the sitting position (Fig 5). The neurologic deficits result from a thrombotic, ischemic inflammation of small blood vessels in the spinal cord and/or brain. The interval between initial EHV-1 infection of the respiratory tract and the subsequent onset of neurological signs is 8 to 12 days. The neurological deficits appear suddenly and reach their peak intensity within 48 hours. The disease is resistant to vaccination, the clinical attack rate is high, and large numbers of fatalities are the rule. The prognosis for non-recumbent horses is favorable, but is poor for animals that remain down for longer than 24 hours.

A major advance in understanding the mechanistic basis of EHV-1 paralytic disease was the recent discovery that a single gene alteration – a mutated version of the viral gene that encodes its replicative polymerase – endows such mutant virus strains with enhanced virulence for the horse. This discovery that a single gene of EHV-1 plays such a prominent role in the viral attribute of hypervirulence has led to a genetic test for identifying latent equine carriers of mutant, neuropathogenic strains of EHV-1.

Figure 5. (video). Neurological manifestation of infection of a horse by a hypervirulent, mutant strain of EHV-1 (neuropathogenic strain). The horse has lost the use of muscles in its hind limbs and is unable to stand.

photo of neurological disease

(click for on image to view video)
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E. EHV-1 Disease in Newborn Foals

Occasionally, foals infected with EHV-1 are born at term but are either obviously sick at birth or become ill within one or two days of parturition. Typically, the clinical progression of the disease is rapid and the fatal outcome unaffected by intensive, supportive veterinary care. The foals fail to nurse, become lethargic, pyrexic, leukopenic, hypoxic, and exhibit severe respiratory distress. Congenital EHV-1 infection can be epidemic in nature and may occur either in association with an outbreak of abortion or without concurrent abortion or obvious respiratory disease in the dams. It is uncertain whether such foals are infected in utero with EHV-1 or acquire a rapid, post-parturient infection from their dams. In either case, viral pneumonia quickly manifests and nearly always leads to respiratory failure and death within a few days (Fig 6).

Figure 6. Full term, newborn foal infected with EHV-1 and exhibiting respiratory distress. Clinical deterioration in congenitally infected newborn foals occurs rapidly, and the prognosis is always grave.

photo of respiratory distress

 

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F. Prevention and Control of EHV-1 Diseases

The two mainstays for minimizing the economic and equine welfare burden caused by infection with EHV-1 are prophylactic vaccination and careful attention to sound herd-management practices.

Vaccination against EHV-1 is recommended as part of the preventive, herd-health program for all horses at risk for acquiring infection. More than a dozen commercially manufactured vaccines (both inactivated and attenuated-live) are currently marketed for use in controlling abortion and respiratory disease caused by EHV-1. No current vaccine has been demonstrated to protect against the neurological manifestation of EHV-1 infection. When used as an adjunct to sound management practices, vaccination can be effective in modulating the severity of EHV-1 respiratory disease in young stock and in limiting the occurrence and severity of abortion storms in broodmares (Fig 7). Annual re-boosting with multiple doses of the vaccines is required for optimal results.

Figure 7. Annual incidence of EHV-1 abortion in Kentucky’s Thoroughbred broodmare population between 1957 and 2003. Widespread vaccination for the disease began in 1981.

graph of incidence of abortion

 

A number of empirically established, herd management procedures have emerged as being effective for reducing the risk of large-scale, farm-wide outbreaks of EHV-1 disease. Management recommendations are based on the practices of (1) subdivision of the at-risk population of horses on the premises into smaller groups; (2) the maintenance of those sub- groups as closed, physically isolated units; and, (3) the minimalization of stress caused to horses by crowding, poor nutritional state, heavy parasite infestation, lengthy transport, disruption of established social groups, inclement weather, en masse weaning, etc. For maximal effectiveness, group size should be as small as the physical facilities will reasonably allow, with each group kept under conditions that limit the transmission of virus between established groups. Restrictions should be placed on movement of horses into and out of each established group, and contact with transient horses should be avoided. The addition of any new horse into a closed group should be preceded by a 21-day period of isolation.

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II. Bibliography

A. Reviews and Overviews

1. van Maanen, C. Equine herpesvirus 1 and 4 infections: an update. Veterinary Quarterly. 2002; 24( 2):58-78; ISSN: 0165-2176. Abstract.

2. Allen, G. P; Kydd, J. H; Slater, J. D, and Smith, K. C. Equid herpesvirus 1 and equid herpesvirus 4 infections. Infectious Diseases of Livestock. 2004; 2(Ed.2):829-859.

3. Reed, S. M and Toribio, R. E. Equine herpesvirus 1 and 4. Veterinary Clinics of North America. Equine Practice. 2004; 20(3):631-642; ISSN: 0749-0739. Abstract.

4. Patel, J. R and Heldens, J. Equine herpesviruses 1 (EHV-1) and 4 (EHV-4)--epidemiology, disease and immunoprophylaxis: a brief review. Veterinary Journal. 2005; 170(1):14-23; ISSN: 1090-0233. Abstract.

5. Smith, K. Equine herpesviruses 1 and 4: creeping to a solution. Veterinary Journal. 2005; 170(1):6-7; ISSN: 1090-0233.

6. Borchers, K.; Thein, R., and Sterner Kock, A. Pathogenesis of equine herpesvirus-associated neurological disease: a revised explanation. Equine Veterinary Journal. 2006; 38(3):283-287; ISSN: 0425-1644.

7. Field, H. J; Biswas, S., and Mohammad, I. T. Herpesvirus latency and therapy--from a veterinary perspective. Antiviral Research. 2006; 71(2-3):127-133; ISSN: 0166-3542. Abstract.

8. Kydd, J. H.; Townsend, H. G., and Hannant, D. The equine immune response to equine herpesvirus-1: the virus and its vaccines. Veterinary Immunology and Immunopathology. 2006; 111(1-2):15-30; ISSN: 0165-2427. Abstract.

9. van der Meulen, K. M.; Favoreel, H. W.; Pensaert, M. B., and Nauwynck, H. J. Immune escape of equine herpesvirus 1 and other herpesviruses of veterinary importance. Veterinary Immunology and Immunopathology. 2006; 111(1-2):31-40; ISSN: 0165-2427. Abstract.

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B. Epidemiology

1. Friday, P. A; Scarratt, W. K; Elvinger, F.; Timoney, P. J, and Bonda, A. Ataxia and paresis with equine herpesvirus type 1 infection in a herd of riding school horses. Journal of Veterinary Internal Medicine. 2000; 14(2):197-201; ISSN: 0891-6640. Abstract.

2. Gilkerson, J. R; Love, D. N, and Whalley, J. M. Incidence of equine herpesvirus 1 infection in thoroughbred weanlings on two stud farms. Australian Veterinary Journal. 2000; 78(4):277-278; ISSN: 0005-0423. Full text.

3. Schroer, U.; Lange, A.; Glatzel, P.; Ludwig, H., and Borchers, K. Die Bedeutung der Infektion mit dem equinen Herpesvirus Typ 1 (EHV-1) in einem deutschen Vollblutgestut: Impfung, Abortgeschehen und Diagnostik [The relevance of equine herpesvirus 1 (EHV-1) infection in a German thoroughbred stud: vaccination, abortion and diagnostics]. Berliner Und Munchener Tierarztliche Wochenschrift. 2000; 113(2):53-59. Abstract.

4. van Maanen, C.; Sloet van Oldruitenborgh Oosterbaan, M. M; Damen, E. A, and Derksen, A. G. Neurological disease associated with EHV-1-infection in a riding school: clinical and virological characteristics. Equine Veterinary Journal. 2001; 33(2):191-196; ISSN: 0425-1644. Abstract.

5. Allen, G. P. Epidemic disease caused by equine herpesvirus-1: recommendations for prevention and control. Equine Veterinary Education (British Edition). 2002; 14(3):136-142; ISSN: 0957-7734.

6. Stierstorfer, B.; Eichhorn, W.; Schmahl, W.; Brandmuller, C.; Kaaden, O. R, and Neubauer, A. Equine herpesvirus type 1 (EHV-1) myeloencephalopathy: a case report. Journal of Veterinary Medicine. B, Infectious Diseases and Veterinary Public Health. 2002; 49(1):37-41; ISSN: 0931-1793. Abstract.

7. Taouji, S.; Collobert, C.; Gicquel, B.; Sailleau, C.; Brisseau, N.; Moussu, C.; Breuil, M. F; Pronost, S.; Borchers, K., and Zientara, S. Detection and isolation of equine herpesviruses 1 and 4 from horses in Normandy: an autopsy study of tissue distribution in relation to vaccination status. Journal of Veterinary Medicine. B, Infectious Diseases and Veterinary Public Health. 2002; 49(8):394-399; ISSN: 0931-1793. Abstract.

8. Allen, G.P.; Powell, D. Equine herpesvirus neurological disease in the USA and United Kingdom. Equine Disease Quarterly. 2003; 12(3):2-3. Full text.

9. Cardwell, J.; Smith, K.; Newton, R.; Blunden, T.; Bestbier, M., and Whitwell, K. EHV paralytic disease in the south of England. Veterinary Record . 2003; 152(14):441-442; ISSN: 0042-4900.

10. Foote, C. E; Gilkerson, J. R; Whalley, J. M, and Love, D. N . Seroprevalence of equine herpesvirus 1 in mares and foals on a large Hunter Valley stud farm in years pre- and postvaccination. Australian Veterinary Journal. 2003; 81(5):283-288; ISSN: 0005-0423. Full text.

11. Studdert, M. J; Hartley, C. A; Dynon, K.; Sandy, J. R; Slocombe, R. F; Charles, J. A; Milne, M. E; Clarke, A. F, and El Hage, C. Outbreak of equine herpesvirus type 1 myeloencephalitis: new insights from virus identification by PCR and the application of an EHV-1-specific antibody detection ELISA. Veterinary Record. 2003; 153(14):417-423; ISSN: 0042-4900. Abstract.

12. Allen, G. P; Kydd, J. H; Slater, J. D, and Smith, K. C. Equid herpesvirus 1 and equid herpesvirus 4 infections. Infectious Diseases of Livestock. 2004; 2(Ed.2):829-859.

13. Foote, C. E.; Love, D. N.; Gilkerson, J. R., and Whalley, J. M. Detection of EHV-1 and EHV-4 DNA in unweaned Thoroughbred foals from vaccinated mares on a large stud farm. Equine Veterinary Journal. 2004; 36(4):341-345; ISSN: 0425-1644. Abstract.

14. Molinkova, D. ; Celer, V. Jr, and Jahn, P. Isolation and partial characterization of equine herpesvirus type 1 in Czechia. Folia Microbiologica. 2004; 49(5):605-611; ISSN: 0015-5632. Abstract.

15. Goehring, L. S; van Maanen, C., and Sloet van Oldruitenborgh Oosterbaan, M. M. Neurological syndromes among horses in The Netherlands. A 5 year retrospective survey (1999-2004). Veterinary Quarterly. 2005; 27(1):11-20; ISSN: 0165-2176. Full text.

16. Patel, J. R and Heldens, J. Equine herpesviruses 1 (EHV-1) and 4 (EHV-4)--epidemiology, disease and immunoprophylaxis: a brief review. Veterinary Journal. 2005; 170(1):14-23; ISSN: 1090-0233. Full text.

17. Bell, S. A.; Balasuriya, U. B.; Gardner, I. A.; Barry, P. A.; Wilson, W. D.; Ferraro, G. L., and MacLachlan, N. J. Temporal detection of equine herpesvirus infections of a cohort of mares and their foals. Veterinary Microbiology. 2006; 116(4):249-257; ISSN: 0378-1135. Abstract.

18. Foote, C. E.; Love, D. N.; Gilkerson, J. R.; Wellington, J. E., and Whalley, J. M. EHV-1 and EHV-4 infection in vaccinated mares and their foals. Veterinary Immunology and Immunopathology. 2006; 111(1-2):41-46; ISSN: 0165-2427. Abstract.

19. Goehring, L. S.; van Winden, S. C.; van Maanen, C., and Sloet van Oldruitenborgh Oosterbaan, M. M. Equine herpesvirus type 1-associated myeloencephalopathy in The Netherlands: a four-year retrospective study (1999-2003). Journal of Veterinary Internal Medicine. 2006; 20(3):601-607; ISSN: 0891-6640. Abstract.

20. Kydd, J. H and Smith, K. C. Equine herpesvirus neurologic disease: reflections from across the pond. Journal of Veterinary Internal Medicine. 2006; 20(3):467-468; ISSN: 0891-6640.

21. Martinez, J. P.; Martin Ocampos, G. P.; Fernandez, L. C.; Fuentealba, N. A.; Cid de la Paz, V.; Barrandeguy, M., and Galosi, C. M. Detection of equine herpesvirus 1 genome 1B in Argentina. Revue Scientifique et Technique. 2006; 25(3):1075-1079; ISSN: 0253-1933. Full text (pdf).

22. Nugent, J.; Birch Machin, I.; Smith, K. C.; Mumford, J. A.; Swann, Z.; Newton, J. R.; Bowden, R. J.; Allen, G. P., and Davis Poynter, N. Analysis of equid herpesvirus 1 strain variation reveals a point mutation of the DNA polymerase strongly associated with neuropathogenic versus nonneuropathogenic disease outbreaks. Journal of Virology. 2006; 80(8):4047-4060; ISSN: 0022-538X. Full text.

23. Henninger, R. W.; Reed, S. M.; Saville, W. J.; Allen, G. P.; Hass, G. F.; Kohn, C. W., and Sofaly, C. Outbreak of neurologic disease caused by equine herpesvirus-1 at a university equestrian center. Journal of Veterinary Internal Medicine. 2007; 21(1):157-165; ISSN: 0891-6640. Abstract.

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C. Pathogenesis

1. Allen, G. A Portrait of the Pathogenesis of Equine Herpesvirus Abortion, Equine Disease Quarterly, October 1998, Vol 7 (#1), p 3-4. Full text.

2. Csellner, H.; Walker, C.; Wellington, J. E; McLure, L. E; Love, D. N, and Whalley, J. M. EHV-1 glycoprotein D (EHV-1 gD) is required for virus entry and cell-cell fusion, and an EHV-1 gD deletion mutant induces a protective immune response in mice. Archives of Virology. 2000; 145(11):2371-2385; ISSN: 0304-8608. Abstract.

3. Del Piero, F. ; Wilkins, P. A; Timoney, P. J; Kadushin, J.; Vogelbacker, H.; Lee, J. W; Berkowitz, S. J, and La Perle, K. M. Fatal nonneurological EHV-1 infection in a yearling filly. Veterinary Pathology. 2000; 37(6):672-676; ISSN: 0300-9858. Full text.

4. Smith, K. C; Whitwell, K. E; Mumford, J. A; Hannant, D.; Blunden, A. S, and Tearle, J. P. Virulence of the V592 isolate of equid herpesvirus-1 in ponies. Journal of Comparative Pathology. 2000; 122(4):288-297; ISSN: 0021-9975. Abstract.

5. Smith, P. M; Zhang, Y.; Grafton, W. D; Jennings, S. R, and O'Callaghan, D. J. Severe murine lung immunopathology elicited by the pathogenic equine herpesvirus 1 strain RacL11 correlates with early production of macrophage inflammatory proteins 1alpha, 1beta, and 2 and tumor necrosis factor alpha. Journal of Virology. 2000; 74(21):10034-10040; ISSN: 0022-538X. Full text.

6. Granzow, H.; Klupp, B. G; Fuchs, W.; Veits, J.; Osterrieder, N., and Mettenleiter, T. C. Egress of alphaherpesviruses: comparative ultrastructural study. Journal of Virology. 2001; 75(8):3675-3684; ISSN: 0022-538X. Full text.

7. Smith, D.; Hamblin, A., and Edington, N. Infection of endothelial cells with equine herpesvirus-1 (EHV-1) occurs where there is activation of putative adhesion molecules: a mechanism for transfer of virus. Equine Veterinary Journal. 2001; 33(2):138-142; ISSN: 0425-1644. Abstract.

8. Smith, K. C and Borchers, K. A study of the pathogenesis of equid herpesvirus-1 (EHV-1) abortion by DNA in-situ hybridization. Journal of Comparative Pathology. 2001; 125(4):304-310; ISSN: 0021-9975. Abstract.

9. Frampton, A. R Jr; Smith, P. M; Zhang, Y.; Matsumura, T.; Osterrieder, N., and O'Callaghan, D. J. Contribution of gene products encoded within the unique short segment of equine herpesvirus 1 to virulence in a murine model. Virus Research. 2002; 90(1-2):287-301; ISSN: 0168-1702. Abstract.

10. Smith, D.; Hamblin, A., and Edington, N. Equid herpesvirus 1 infection of endothelial cells requires activation of putative adhesion molecules: an in vitro model. Clinical and Experimental Immunology. 2002; 129(2):281-287; ISSN: 0009-9104. Abstract.

11. Taouji, S.; Collobert, C.; Gicquel, B.; Sailleau, C.; Brisseau, N.; Moussu, C.; Breuil, M. F; Pronost, S.; Borchers, K., and Zientara, S. Detection and isolation of equine herpesviruses 1 and 4 from horses in Normandy: an autopsy study of tissue distribution in relation to vaccination status. Journal of Veterinary Medicine. B, Infectious Diseases and Veterinary Public Health. 2002; 49(8):394-399; ISSN: 0931-1793. Abstract.

12. Albrecht, R. A; Jang, H. K; Kim, S. K, and O'Callaghan, D. J. Direct interaction of TFIIB and the IE protein of equine herpesvirus 1 is required for maximal trans-activation function. Virology. 2003; 316(2):302-312; ISSN: 0042-6822. Abstract.

13. Kirisawa, R. ; Kobayashi, T.; Uematsu, R.; Ikeda, A.; Kuroiwa, R.; Urakami, A., and Iwai, H. Growth of recombinant equine herpesvirus 1 (EHV-1) replaced with passage-induced mutant gene 1 and gene 71 derived from an attenuated EHV-1 in cell cultures and in the lungs of mice. Veterinary Microbiology. 2003; 95(3):159-174; ISSN: 0378-1135. Abstract.

14. Rappocciolo, G.; Birch, J., and Ellis, S. A. Down-regulation of MHC class I expression by equine herpesvirus-1. Journal of General Virology. 2003; 84(Pt 2):293-300; ISSN: 0022-1317. Full text.

15. Szeredi, L.; Palfi, V., and Molnar, T. Comparison of methods for the diagnosis of equine herpesvirus type 1 infection. Acta Veterinaria Hungarica. 2003; 51(2):153-163; ISSN: 0236-6290. Abstract.

16. Tearle, J. P ; Smith, K. C; Platt, A. J; Hannant, D.; Davis Poynter, N. J, and Mumford, J. A. In vitro characterisation of high and low virulence isolates of equine herpesvirus-1 and -4. Research in Veterinary Science. 2003; 75(1):83-86; ISSN: 0034-5288. Abstract.

17. Zhang, Y.; Smith, P. M; Frampton, A. R; Osterrieder, N.; Jennings, S. R, and O'Callaghan, D. J. Cytokine profiles and long-term virus-specific antibodies following immunization of CBA mice with equine herpesvirus 1 and viral glycoprotein D. Viral Immunology. 2003; 16(3):307-320; ISSN: 0882-8245. Abstract.

18. Allen, G. Equine rhinopneumonitis. In. OIE Manual of Diagnostic Tests and Vaccines for Terrestrial Animals. 5th ed. Paris: OIE; 2004;(Chapt. 2.5.7): pp. 707-716. Full text.

19. Ibrahim, el S. M.; Pagmajav, O.; Yamaguchi, T.; Matsumura, T., and Fukushi, H. Growth and virulence alterations of equine herpesvirus 1 by insertion of a green fluorescent protein gene in the intergenic region between ORFs 62 and 63. Microbiology and Immunology. 2004; 48(11):831-842; ISSN: 0385-5600. Full text.

20. Neubauer, A. and Osterrieder, N. Equine herpesvirus type 1 (EHV-1) glycoprotein K is required for efficient cell-to-cell spread and virus egress. Virology. 2004; 329(1):18-32; ISSN: 0042-6822. Abstract.

21. von Einem, J.; Wellington, J.; Whalley, J. M; Osterrieder, K.; O'Callaghan, D. J, and Osterrieder, N. The truncated form of glycoprotein gp2 of equine herpesvirus 1 (EHV-1) vaccine strain KyA is not functionally equivalent to full-length gp2 encoded by EHV-1 wild-type strain RacL11. Journal of Virology. 2004; 78( 6):3003-3013; ISSN: 0022-538X. Full text.

22. Frampton, A. R Jr; Goins, W. F; Cohen, J. B; von Einem, J. ; Osterrieder, N.; O'Callaghan, D. J, and Glorioso, J. C. Equine herpesvirus 1 utilizes a novel herpesvirus entry receptor. Journal of Virology. 2005; 79(5):3169-3173; ISSN: 0022-538X. Full text.

23. Goehring, L. S; van Maanen, C., and Sloet van Oldruitenborgh Oosterbaan, M. M. Neurological syndromes among horses in The Netherlands. A 5 year retrospective survey (1999-2004). Veterinary Quarterly. 2005; 27(1):11-20; ISSN: 0165-2176. Full text.

24. Pagamjav, O. ; Sakata, T.; Matsumura, T.; Yamaguchi, T., and Fukushi, H. Natural recombinant between equine herpesviruses 1 and 4 in the ICP4 gene. Microbiology and Immunology. 2005; 49(2):167-179; ISSN: 0385-5600. Full text.

25. Smith, P. M; Kahan, S. M; Rorex, C. B; von Einem, J.; Osterrieder, N., and O'Callaghan, D. J. Expression of the full-length form of gp2 of equine herpesvirus 1 (EHV-1) completely restores respiratory virulence to the attenuated EHV-1 strain KyA in CBA mice. Journal of Virology. 2005; 79(8):5105-5115; ISSN: 0022-538X. Full text.

26. Allen, G. P. and Breathnach, C. C. Quantification by real-time PCR of the magnitude and duration of leucocyte-associated viraemia in horses infected with neuropathogenic vs. non-neuropathogenic strains of EHV-1. Equine Veterinary Journal. 2006; 38(3):252-257; ISSN: 0425-1644. Abstract.

27. Borchers, K.; Thein, R., and Sterner Kock, A. Pathogenesis of equine herpesvirus-associated neurological disease: a revised explanation. Equine Veterinary Journal. 2006; 38(3):283-287; ISSN: 0425-1644.

28. Chiam, R.; Smid, L.; Kydd, J. H.; Smith, K. C.; Platt, A., and  Davis Poynter, N. J. Use of polarised equine endothelial cell cultures and an in vitro thrombosis model for potential characterisation of EHV-1 strain variation. Veterinary Microbiology. 2006; 113(3-4):243-249; ISSN: 0378-1135. Abstract.

29. Galosi, C. M.; Barbeito, C. G.; Martin Ocampos, G. P.; Martinez, J. P.; Ayala, M. A.; Corva, S. G.; Fuentealba, N. A., and Gimeno, E. J. An Argentine equine herpesvirus strain with special restriction patterns protect mice challenged with a pathogenic strain. Journal of Veterinary Medicine. B. Infectious Diseases and Veterinary Public Health. 2006; 53(9):412-417; ISSN: 0931-1793. Abstract.

30. Nugent, J.; Birch Machin, I.; Smith, K. C.; Mumford, J. A.; Swann, Z.; Newton, J. R.; Bowden, R. J.; Allen, G. P., and Davis Poynter, N. Analysis of equid herpesvirus 1 strain variation reveals a point mutation of the DNA polymerase strongly associated with neuropathogenic versus nonneuropathogenic disease outbreaks. Journal of Virology. 2006; 80(8):4047-4060; ISSN: 0022-538X. Full text.

31. von Einem, J.; Schumacher, D.; O'Callaghan, D. J., and Osterrieder, N. The alpha-TIF (VP16) homologue (ETIF) of equine herpesvirus 1 is essential for secondary envelopment and virus egress. Journal of Virology. 2006; 80(6):2609-2620; ISSN: 0022-538X. Full text.

32. Ibrahim, E. S.; Kinoh, M.; Matsumura, T.; Kennedy, M.; Allen, G. P.; Yamaguchi, T., and Fukushi, H. Genetic relatedness and pathogenicity of equine herpesvirus 1 isolated from onager, zebra and gazelle.  Archives of Virology. 2007; 152(2):245-255; ISSN: 0304-8608. Abstract.

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D. EHV-1 Respiratory Disease

1. Del Piero, F. ; Wilkins, P. A; Timoney, P. J; Kadushin, J.; Vogelbacker, H.; Lee, J. W; Berkowitz, S. J, and La Perle, K. M. Fatal nonneurological EHV-1 infection in a yearling filly. Veterinary Pathology. 2000; 37(6):672-676; ISSN: 0300-9858. Full text.

2. van Maanen, C.; Vreeswijk, J.; Moonen, P.; Brinkhof, J. ; de Boer Luijtze, E., and Terpstra, C. Differentiation and genomic and antigenic variation among fetal, respiratory, and neurological isolates from EHV1 and EHV4 infections in The Netherlands. Veterinary Quarterly. 2000; 22(2):88-93; ISSN: 0165-2176. Abstract.

3. Del Piero, F. and Wilkins, P. A. Pulmonary vasculotropic EHV-1 infection in equids. Veterinary Pathology. 2001; 38(4):474; ISSN: 0300-9858. Full text.

4. Varrasso, A.; Dynon, K.; Ficorilli, N.; Hartley, C. A; Studdert, M. J, and Drummer, H. E. Identification of equine herpesviruses 1 and 4 by polymerase chain reaction. Australian Veterinary Journal. 2001; 79(8):563-569; ISSN: 0005-0423. Full text.

5. Allen, G. “Respiratory infections by equine herpesvirus types 1 and 4,” In: Lekeux, P. editor, Equine respiratory diseases. International Veterinary Information Service; 2002. Full text.

6. van Maanen, C. Equine herpesvirus 1 and 4 infections: an update. Veterinary Quarterly. 2002; 24(2):58-78; ISSN: 0165-2176. Abstract.

7. Mumford, E. L ; Traub Dargatz, J. L; Carman, J.; Callan, R. J; Collins, J. K; Goltz, K. L; Romm, S. R; Tarr, S. F, and Salman, M. D. Occurrence of infectious upper respiratory tract disease and response to vaccination in horses on six sentinel premises in northern Colorado. Equine Veterinary Journal. 2003; 35(1):72-77; ISSN: 0425-1644. Abstract.

8. Patel, J. R; Foldi, J.; Bateman, H.; Williams, J.; Didlick, S., and Stark, R. Equid herpesvirus (EHV-1) live vaccine strain C147: efficacy against respiratory diseases following EHV types 1 and 4 challenges. Veterinary Microbiology. 2003; 92(1-2):1-17; ISSN: 0378-1135. Abstract.

9. Allen, G. Equine rhinopneumonitis. In. OIE Manual of Diagnostic Tests and Vaccines for Terrestrial Animals. 5th ed. Paris: OIE; 2004;(Chapt. 2.5.7): pp. 707-716. Full text.

10. Maeda, K.; Kai, K.; Hayashi, T.; Hasegawa, K., and Matsumura, T. Intercellular adhesion molecule-1 (ICAM-1) and lymphocyte function-associated antigen-1 (LFA-1) contribute to the elimination of equine herpesvirus type 1 (EHV-1) from the lungs of intranasally infected BALB/c mice. Journal of Comparative Pathology. 2004; 130(2-3):162-170; ISSN: 0021-9975 . Abstract.

11. Smith, P. M.; Kahan, S. M.; Rorex, C. B.; von Einem, J.; Osterrieder, N., and O'Callaghan, D. J. Expression of the full-length form of gp2 of equine herpesvirus 1 (EHV-1) completely restores respiratory virulence to the attenuated EHV-1 strain KyA in CBA mice. Journal of Virology. 2005; 79(8):5105-5115; ISSN: 0022-538X. Full text.

12. Breathnach, C. C.; Yeargan, M. R.; Timoney, J. F., and Allen, G. P. Detection of equine herpesvirus-specific effector and memory cytotoxic immunity in the equine upper respiratory tract. Veterinary Immunology and Immunopathology. 2006; 111(1-2):117-125; ISSN: 0165-2427. Abstract.

13. Coombs, D. K.; Patton, T.; Kohler, A. K.; Soboll, G.; Breathnach, C.; Townsend, H. G., and Lunn, D. P. Cytokine responses to EHV-1 infection in immune and non-immune ponies. Veterinary Immunology and Immunopathology. 2006; 111(1-2):109-116; ISSN: 0165-2427. Abstract.

14. Foote, C. E.; Raidal, S. L.; Pecenpetelovska, G.; Wellington, J. E., and Whalley, J. M. Inoculation of mares and very young foals with EHV-1 glycoproteins D and B reduces virus shedding following respiratory challenge with EHV-1. Veterinary Immunology and Immunopathology. 2006; 111(1-2):97-108; ISSN: 0165-2427. Abstract.

15. Hussey, S. B.; Clark, R.; Lunn, K. F.; Breathnach, C.; Soboll, G.; Whalley, J. M., and Lunn, D. P. Detection and quantification of equine herpesvirus-1 viremia and nasal shedding by real-time polymerase chain reaction. Journal of Veterinary Diagnostic Investigation. 2006; 18(4):335-342; ISSN: 1040-6387. Abstract.

16. Dynon, K.; Black, W. D.; Ficorilli, N.; Hartley, C. A., and Studdert, M. J. Detection of viruses in nasal swab samples from horses with acute, febrile, respiratory disease using virus isolation, polymerase chain reaction and serology. Australian Veterinary Journal. 2007; 85(1-2):46-50; ISSN: 0005-0423. Abstract.

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E. EHV-1 Abortion

1. Allen, G. A portrait of the pathogenesis of equine herpesvirus abortion. Equine Disease Quarterly. October 1998; 7(1):3-4. Full text.

2. Mukaiya, R.; Kimura, T.; Ochiai, K.; Wada, R., and Umemura, T. Demonstration of equine herpesvirus-1 gene expression in the placental trophoblasts of naturally aborted equine fetuses. Journal of Comparative Pathology. 2000; 123(2-3):119-125; ISSN: 0021-9975. Abstract.

3. Schroer, U.; Lange, A.; Glatzel, P.; Ludwig, H., and Borchers, K. Die Bedeutung der Infektion mit dem equinen Herpesvirus Typ 1 (EHV-1) in einem deutschen Vollblutgestut: Impfung, Abortgeschehen und Diagnostik [The relevance of equine herpesvirus 1 (EHV-1) infection in a German thoroughbred stud: vaccination, abortion and diagnostics]. Berliner Und Munchener Tierarztliche Wochenschrift. 2000; 113(2):53-59. Abstract.

4. Smith, K. C; Whitwell, K. E; Mumford, J. A; Hannant, D.; Blunden, A. S, and Tearle, J. P. Virulence of the V592 isolate of equid herpesvirus-1 in ponies. Journal of Comparative Pathology. 2000; 122(4):288-297; ISSN: 0021-9975. Abstract.

5. van Maanen, C.; Vreeswijk, J.; Moonen, P.; Brinkhof, J. ; de Boer Luijtze, E., and Terpstra, C. Differentiation and genomic and antigenic variation among fetal, respiratory, and neurological isolates from EHV1 and EHV4 infections in The Netherlands. Veterinary Quarterly. 2000; 22(2):88-93; ISSN: 0165-2176. Abstract.

6. van Maanen, C.; Willink, D. L; Smeenk, L. A; Brinkhof, J., and Terpstra, C. An equine herpesvirus 1 (EHV1) abortion storm at a riding school. Veterinary Quarterly. 2000; 22(2):83-87; ISSN: 0165-2176. Abstract.

7. Walker, C.; Ruitenberg, K. M; Love, D. N, and Millar Whalley, J. Immunization of BALB/c mice with DNA encoding equine herpesvirus 1 (EHV-1) glycoprotein D affords partial protection in a model of EHV-1-induced abortion. Veterinary Microbiology. 2000; 76(3):211-220; ISSN: 0378-1135. Abstract.

8. Galosi, C. M; Vila Roza, M. V; Oliva, G. A; Pecoraro, M. R; Echeverria, M. G; Corva, S., and Etcheverrigaray, M. E. A polymerase chain reaction for detection of equine herpesvirus-1 in routine diagnostic submissions of tissues from aborted foetuses. Journal of Veterinary Medicine. B, Infectious Diseases and Veterinary Public Health. 2001; 48(5):341-346; ISSN: 0931-1793. Abstract.

9. Smith, K. C and Borchers, K. A study of the pathogenesis of equid herpesvirus-1 (EHV-1) abortion by DNA in-situ hybridization. Journal of Comparative Pathology. 2001; 125(4):304-310; ISSN: 0021-9975. Abstract.

10. Varrasso, A. ; Dynon, K.; Ficorilli, N.; Hartley, C. A; Studdert, M. J, and Drummer, H. E. Identification of equine herpesviruses 1 and 4 by polymerase chain reaction. Australian Veterinary Journal. 2001; 79(8):563-569; ISSN: 0005-0423. Full text.

11. Iqbal, J. and Edington, N. Equid herpesvirus 1 is neurotropic in mice, but latency from which infectious virus can be reactivated does not occur. Acta Veterinaria Hungarica. 2002; 50(1):117-129; ISSN: 0236-6290. Abstract.

12. van Maanen, C. Equine herpesvirus 1 and 4 infections: an update. Veterinary Quarterly. 2002; 24(2):58-78; ISSN: 0165-2176. Abstract.

13. Daly, P. and Doyle, S. The development of a competitive PCR-ELISA for the detection of equine herpesvirus-1. Journal of Virological Methods . 2003; 107(2):237-244; ISSN: 0166-0934. Abstract.

14. Gerst, S.; Borchers, K.; Gower, S. M, and Smith, K. C. Detection of EHV-1 and EHV-4 in placental sections of naturally occurring EHV-1- and EHV-4-related abortions in the UK: use of the placenta in diagnosis. Equine Veterinary Journal. 2003; 35(5):430-433; ISSN: 0425-1644. Abstract.

15. Kydd, J. H; Wattrang, E., and Hannant, D. Pre-infection frequencies of equine herpesvirus-1 specific, cytotoxic T lymphocytes correlate with protection against abortion following experimental infection of pregnant mares. Veterinary Immunology and Immunopathology. 2003; 96(3-4):207-217; ISSN: 0165-2427. Abstract.

16. Patel, J. R; Bateman, H.; Williams, J., and Didlick, S. Derivation and characterisation of a live equid herpes virus-1 (EHV-1) vaccine to protect against abortion and respiratory disease due to EHV-1. Veterinary Microbiology. 2003; 91(1):23-39; ISSN: 0378-1135. Abstract.

17. Studdert, M. J; Hartley, C. A; Dynon, K.; Sandy, J. R; Slocombe, R. F; Charles, J. A; Milne, M. E; Clarke, A. F, and El Hage, C. Outbreak of equine herpesvirus type 1 myeloencephalitis: new insights from virus identification by PCR and the application of an EHV-1-specific antibody detection ELISA. Veterinary Record. 2003; 153(14):417-423; ISSN: 0042-4900. Abstract.

18. Szeredi, L.; Aupperle, H., and Steiger, K. Detection of equine herpesvirus-1 in the fetal membranes of aborted equine fetuses by immunohistochemical and in-situ hybridization techniques. Journal of Comparative Pathology. 2003; 129(2-3):147-153; ISSN: 0021-9975. Abstract.

19. Szeredi, L.; Palfi, V., and Molnar, T. Comparison of methods for the diagnosis of equine herpesvirus type 1 infection. Acta Veterinaria Hungarica. 2003; 51(2):153-163; ISSN: 0236-6290. Abstract.

20. Allen, G. Equine rhinopneumonitis. In. OIE Manual of Diagnostic Tests and Vaccines for Terrestrial Animals. 5th ed. Paris: OIE; 2004;(Chapt. 2.5.7): pp. 707-716. Full text.

21. Galosi, C. M; Barbeito, C. G; Vila Roza, M. V; Cid de la Paz, V.; Ayala, M. A; Corva, S. G; Etcheverrigaray, M. E, and Gimeno, E. J. Argentine strain of equine herpesvirus 1 isolated from an aborted foetus shows low virulence in mouse respiratory and abortion models. Veterinary Microbiology. 2004; 103(1-2 ):1-12; ISSN: 0378-1135. Abstract.

22. Kimura, T.; Hasebe, R.; Mukaiya, R.; Ochiai, K.; Wada, R., and Umemura, T. Decreased expression of equine herpesvirus-1 early and late genes in the placenta of naturally aborted equine fetuses. Journal of Comparative Pathology. 2004; 130(1):41-47; ISSN: 0021-9975. Abstract.

23. Kondo, T.; McGregor, M.; Chu, Q.; Chen, D.; Horimoto, T., and Kawaoka, Y. A protective effect of epidermal powder immunization in a mouse model of equine herpesvirus-1 infection. Virology. 2004; 318(1):414-419; ISSN: 0042-6822. Abstract.

24. Reed, S. M and Toribio, R. E. Equine herpesvirus 1 and 4. Veterinary Clinics of North America. Equine Practice. 2004; 20(3):631-642; ISSN: 0749-0739. Abstract.

25. Smith, K. C; Whitwell, K. E; Blunden, A. S; Bestbier, M. E ; Scase, T. J; Geraghty, R. J; Nugent, J.; Davis Poynter, N. J, and Cardwell, J. M. Equine herpesvirus-1 abortion: atypical cases with lesions largely or wholly restricted to the placenta. Equine Veterinary Journal. 2004; 36(1):79-82; ISSN: 0425-1644.

26. Gupta, A. K.; Kaur, D.; Rattan, B., and Yadav, M. P. Molecular variability in different Indian isolates of equine herpesvirus-1. Veterinary Research Communications. 2005; 29(8):721-734; ISSN: 0165-7380. Abstract.

27. Elia, G.; Decaro, N.; Martella, V.; Campolo, M.; Desario, C.; Lorusso, E.; Cirone, F., and Buonavoglia, C. Detection of equine herpesvirus type 1 by real time PCR. Journal of Virological Methods. 2006; 133(1):70-75; ISSN: 0166-0934. Abstract.

28. Martinez, J. P.; Martin Ocampos, G. P.; Fernandez, L. C.; Fuentealba, N. A.; Cid de la Paz, V.; Barrandeguy, M., and Galosi, C. M. Detection of equine herpesvirus 1 genome 1B in Argentina. Revue Scientifique et Technique. 2006; 25(3):1075-1079; ISSN: 0253-1933. Full text (pdf).

29. Paillot, R.; Daly, J. M.; Luce, R.; Montesso, F.; Davis Poynter, N.; Hannant, D., and Kydd, J. H. Frequency and phenotype of EHV-1 specific, IFN-gamma synthesising lymphocytes in ponies: the effects of age, pregnancy and infection. Developmental and Comparative Immunology. 2007; 31(2):202-214; ISSN: 0145-305X. Abstract.

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F. EHV-1 Neurological Disease

1. Friday, P. A; Scarratt, W. K; Elvinger, F.; Timoney, P. J, and Bonda, A. Ataxia and paresis with equine herpesvirus type 1 infection in a herd of riding school horses. Journal of Veterinary Internal Medicine. 2000; 14(2):197-201; ISSN: 0891-6640. Abstract.

2. van Maanen, C.; Vreeswijk, J.; Moonen, P.; Brinkhof, J. ; de Boer Luijtze, E., and Terpstra, C. Differentiation and genomic and antigenic variation among fetal, respiratory, and neurological isolates from EHV1 and EHV4 infections in The Netherlands. Veterinary Quarterly. 2000; 22(2):88-93; ISSN: 0165-2176. Abstract.

3. Olsen, T. F. Equine herpesvirus myeloencephalopathy in a 14-year-old quarter horse stallion. Canadian Veterinary Journal. 2001; 42(3):217-220; ISSN: 0008-5286. Abstract.

4. van Maanen, C.; Sloet van Oldruitenborgh Oosterbaan, M. M; Damen, E. A, and Derksen, A. G. Neurological disease associated with EHV-1-infection in a riding school: clinical and virological characteristics. Equine Veterinary Journal. 2001; 33(2):191-196; ISSN: 0425-1644. Abstract.

5. Hasebe, R.; Kimura, T.; Nakamura, K.; Okazaki, K.; Ochiai, K.; Wada, R., and Umemura, T. Passage of equine herpesvirus-1 in suckling mouse brain enhances extraneural virus growth and subsequent hematogenous neuroinvasion. Journal of Veterinary Medical Science. 2002; 64(10):907-912; ISSN: 0916-7250. Full text.

6. Hasebe, R.; Kimura, T.; Sato, E.; Okazaki, K.; Ochiai, K.; Wada, R., and Umemura, T. Equine herpesvirus-1-induced encephalomyelitis in mice: a comparative study of neuroadapted virus and its parental strain. Journal of Comparative Pathology. 2002; 127(2-3 ):118-125; ISSN: 0021-9975. Abstract.

7. Stierstorfer, B.; Eichhorn, W.; Schmahl, W.; Brandmuller, C.; Kaaden, O. R, and Neubauer, A. Equine herpesvirus type 1 (EHV-1) myeloencephalopathy: a case report. Journal of Veterinary Medicine. B, Infectious Diseases and Veterinary Public Health. 2002; 49(1):37-41; ISSN: 0931-1793. Abstract.

8. van Maanen, C. Equine herpesvirus 1 and 4 infections: an update. Veterinary Quarterly. 2002; 24(2):58-78; ISSN: 0165-2176. Abstract.

9. Allen, G. and Powell, D., “Equine Herpesvirus Neurological Disease in the USA and United Kingdom,” Equine Disease Quarterly, July 2003, Vol 12 (#3), p 2-3. Full text.

10. Cardwell, J.; Smith, K.; Newton, R.; Blunden, T.; Bestbier, M., and Whitwell, K. EHV paralytic disease in the south of England. Veterinary Record . 2003; 152(14):441-442; ISSN: 0042-4900.

11. Studdert, M. J; Hartley, C. A; Dynon, K.; Sandy, J. R; Slocombe, R. F; Charles, J. A; Milne, M. E; Clarke, A. F, and El Hage, C. Outbreak of equine herpesvirus type 1 myeloencephalitis: new insights from virus identification by PCR and the application of an EHV-1-specific antibody detection ELISA. Veterinary Record. 2003; 153(14):417-423; ISSN: 0042-4900. Abstract.

12. Allen, G. P; Kydd, J. H; Slater, J. D, and Smith, K. C. Equid herpesvirus 1 and equid herpesvirus 4 infections. Infectious Diseases of Livestock. 2004; 2(Ed.2):829-859.

13. Chvala, S.; Nowotny, N.; Kotzab, E.; Cain, M., and van den Hoven, R. Use of the meridian test for the detection of equine herpesvirus type 1 infection in horses with decreased performance. Journal of the American Veterinary Medical Association. 2004; 225(4):554-559; ISSN: 0003-1488. Abstract.

14. Frampton, A. R Jr; Smith, P. M; Zhang, Y.; Grafton, W. D; Matsumura, T.; Osterrieder, N., and O'Callaghan, D. J. Meningoencephalitis in mice infected with an equine herpesvirus 1 strain KyA recombinant expressing glycoprotein I and glycoprotein E. Virus Genes. 2004; 29(1):9-17; ISSN: 0920-8569. Abstract.

15. Goehring, L. S; van Maanen, C., and Sloet van Oldruitenborgh Oosterbaan, M. M. Neurological syndromes among horses in The Netherlands. A 5 year retrospective survey (1999-2004). Veterinary Quarterly. 2005; 27(1):11-20; ISSN: 0165-2176. Full text.

16. Sloet van Oldruitenborgh Oosterbaan, M. Neurologische vorm van rhinopneumonie. [Neurologic form of rhinopneumonia]. Tijdschrift Voor Diergeneeskunde. 2005; 130(20):629-631; ISSN: 0040-7453.

17. Allen, G. P. Antemortem detection of latent infection with neuropathogenic strains of equine herpesvirus-1 in horses.  American Journal of Veterinary Research. 2006; 67(8):1401-1405; ISSN: 0002-9645. Abstract.

18. Allen, G. P. and Breathnach, C. C. Quantification by real-time PCR of the magnitude and duration of leucocyte-associated viraemia in horses infected with neuropathogenic vs. non-neuropathogenic strains of EHV-1. Equine Veterinary Journal. 2006; 38(3):252-257; ISSN: 0425-1644. Abstract.

19. Borchers, K.; Thein, R., and Sterner Kock, A. Pathogenesis of equine herpesvirus-associated neurological disease: a revised explanation. Equine Veterinary Journal. 2006; 38(3):283-287; ISSN: 0425-1644.

20. Goehring, L. S.; van Winden, S. C.; van Maanen, C., and Sloet van Oldruitenborgh Oosterbaan, M. M. Equine herpesvirus type 1-associated myeloencephalopathy in The Netherlands: a four-year retrospective study (1999-2003). Journal of Veterinary Internal Medicine. 2006; 20(3):601-607; ISSN: 0891-6640. Abstract.

21. Goodman, L. B.; Wagner, B.; Flaminio, M. J.; Sussman, K. H.; Metzger, S. M.; Holland, R., and Osterrieder, N. Comparison of the efficacy of inactivated combination and modified-live virus vaccines against challenge infection with neuropathogenic equine herpesvirus type 1 (EHV-1). Vaccine. 2006; 24(17):3636-3645; ISSN: 0264-410X. Abstract.

22. Hasebe, R.; Kimura, T.; Nakamura, K.; Ochiai, K.; Okazaki, K.; Wada, R., and Umemura, T. Differential susceptibility of equine and mouse brain microvascular endothelial cells to equine herpesvirus 1 infection. Archives of Virology. 2006; 151(4):775-786; ISSN: 0304-8608. Abstract.

23. Nugent, J.; Birch Machin, I.; Smith, K. C.; Mumford, J. A.; Swann, Z.; Newton, J. R.; Bowden, R. J.; Allen, G. P., and Davis Poynter, N. Analysis of equid herpesvirus 1 strain variation reveals a point mutation of the DNA polymerase strongly associated with neuropathogenic versus nonneuropathogenic disease outbreaks. Journal of Virology. 2006; 80(8):4047-4060; ISSN: 0022-538X. Full text.

24. Pusterla, N.; Wilson, W. D.; Conrad, P. A.; Barr, B. C.; Ferraro, G. L.; Daft, B. M., and Leutenegger, C. M. Cytokine gene signatures in neural tissue of horses with equine protozoal myeloencephalitis or equine herpes type 1 myeloencephalopathy. Veterinary Record. 2006; 159(11):341-346; ISSN: 0042-4900. Abstract.

25. Henninger, R. W.; Reed, S. M.; Saville, W. J.; Allen, G. P.; Hass, G. F.; Kohn, C. W., and Sofaly, C. Outbreak of neurologic disease caused by equine herpesvirus-1 at a university equestrian center. Journal of Veterinary Internal Medicine. 2007; 21(1):157-165; ISSN: 0891-6640. Abstract.

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G. EHV-1 Infection of Newborn Foals

1. Carvalho, R.; Oliveira, A. M; Souza, A. M; Passos, L. M, and Martins, A. S. Prevalence of equine herpesvirus type 1 latency detected by polymerase chain reaction. Archives of Virology. 2000; 145(9):1773-1787; ISSN: 0304-8608. Abstract.

2. van Maanen, C.; Vreeswijk, J.; Moonen, P.; Brinkhof, J. ; de Boer Luijtze, E., and Terpstra, C. Differentiation and genomic and antigenic variation among fetal, respiratory, and neurological isolates from EHV1 and EHV4 infections in The Netherlands. Veterinary Quarterly. 2000; 22(2):88-93; ISSN: 0165-2176. Abstract.

3. Foote, C. E; Gilkerson, J. R; Whalley, J. M, and Love, D. N . Seroprevalence of equine herpesvirus 1 in mares and foals on a large Hunter Valley stud farm in years pre- and postvaccination. Australian Veterinary Journal. 2003; 81(5):283-288; ISSN: 0005-0423. Full text.

4. Szeredi, L.; Palfi, V., and Molnar, T. Comparison of methods for the diagnosis of equine herpesvirus type 1 infection. Acta Veterinaria Hungarica. 2003; 51(2):153-163; ISSN: 0236-6290. Abstract.

5. Foote, C. E; Love, D. N; Gilkerson, J. R, and Whalley, J. M . Detection of EHV-1 and EHV-4 DNA in unweaned Thoroughbred foals from vaccinated mares on a large stud farm. Equine Veterinary Journal. 2004; 36(4):341-345; ISSN: 0425-1644. Abstract.

6. Patel, J. R; Didlick, S., and Bateman, H. Efficacy of a live equine herpesvirus-1 (EHV-1) strain C147 vaccine in foals with maternally-derived antibody: protection against EHV-1 infection. Equine Veterinary Journal. 2004; 36(5):447-451; ISSN: 0425-1644. Abstract.

7. Foote, C. E; Love, D. N; Gilkerson, J. R; Rota, J.; Trevor Jones, P.; Ruitenberg, K. M; Wellington, J. E, and Whalley, J. M. Serum antibody responses to equine herpesvirus 1 glycoprotein D in horses, pregnant mares and young foals. Veterinary Immunology and Immunopathology. 2005; 105(1-2):47-57; ISSN: 0165-2427. Abstract.

8. Kirisawa, R.; Hosoi, Y.; Yamaya, R.; Taniyama, H.; Okamoto, M.; Tsunoda, N.; Hagiwara, K., and Iwai, H. Isolation of equine herpesvirus-1 lacking glycoprotein C from a dead neonatal foal in Japan. Archives of Virology. 2005; 150(12):2549-2565; ISSN: 0304-8608. Abstract.

9. Bell, S. A.; Balasuriya, U. B.; Gardner, I. A.; Barry, P. A.; Wilson, W. D.; Ferraro, G. L., and MacLachlan, N. J. Temporal detection of equine herpesvirus infections of a cohort of mares and their foals. Veterinary Microbiology. 2006; 116(4):249-257; ISSN: 0378-1135. Abstract.

10. Foote, C. E.; Love, D. N.; Gilkerson, J. R.; Wellington, J. E., and Whalley, J. M. EHV-1 and EHV-4 infection in vaccinated mares and their foals. Veterinary Immunology and Immunopathology. 2006; 111(1-2):41-46; ISSN: 0165-2427. Abstract.

11. Foote, C. E.; Raidal, S. L.; Pecenpetelovska, G.; Wellington, J. E., and Whalley, J. M. Inoculation of mares and very young foals with EHV-1 glycoproteins D and B reduces virus shedding following respiratory challenge with EHV-1. Veterinary Immunology and Immunopathology. 2006; 111(1-2):97-108; ISSN: 0165-2427. Abstract.

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H. Prevention and Control

1. Alber, D. G; Killington, R. A, and Stokes, A. Solid matrix-antibody-antigen complexes incorporating equine herpesvirus 1 glycoproteins C and D elicit anti-viral immune responses in BALB/c (H-2K(d)) and C3H (H-2K(k)) mice. Vaccine. 2000; 19(7-8):895-901; ISSN: 0264-410X. Abstract.

2. Birch Machin, I.; Ryder, S.; Taylor, L.; Iniguez, P.; Marault, M.; Ceglie, L.; Zientara, S.; Cruciere, C.; Cancellotti, F.; Koptopoulos, G.; Mumford, J.; Binns, M.; Davis Poynter, N., and Hannant, D. Utilisation of bacteriophage display libraries to identify peptide sequences recognised by equine herpesvirus type 1 specific equine sera. Journal of Virological Methods. 2000; 88(1):89-104; ISSN: 0166-0934. Abstract.

3. Csellner, H.; Walker, C.; Wellington, J. E; McLure, L. E; Love, D. N, and Whalley, J. M. EHV-1 glycoprotein D (EHV-1 gD) is required for virus entry and cell-cell fusion, and an EHV-1 gD deletion mutant induces a protective immune response in mice. Archives of Virology. 2000; 145(11):2371-2385; ISSN: 0304-8608. Abstract.

4. Ruitenberg, K. M; Love, D. N; Gilkerson, J. R; Wellington, J. E, and Whalley, J. M. Equine herpesvirus 1 (EHV-1) glycoprotein D DNA inoculation in horses with pre-existing EHV-1/EHV-4 antibody. Veterinary Microbiology. 2000; 76(2):117-127; ISSN: 0378-1135. Abstract.

5. Ruitenberg, K. M; Walker, C.; Love, D. N; Wellington, J. E, and Whalley, J. M. A prime-boost immunization strategy with DNA and recombinant baculovirus-expressed protein enhances protective immunogenicity of glycoprotein D of equine herpesvirus 1 in naive and infection-primed mice. Vaccine. 2000; 18(14):1367-1373; ISSN: 0264-410X. Abstract.

6. Schroer, U.; Lange, A.; Glatzel, P.; Ludwig, H., and Borchers, K. Die Bedeutung der Infektion mit dem equinen Herpesvirus Typ 1 (EHV-1) in einem deutschen Vollblutgestut: Impfung, Abortgeschehen und Diagnostik [The relevance of equine herpesvirus 1 (EHV-1) infection in a German thoroughbred stud: vaccination, abortion and diagnostics]. Berliner Und Munchener Tierarztliche Wochenschrift. 2000; 113(2):53-59. Abstract.

7. Skinner, G. R and Davies, J. Efficacy of an inactivated vaccine for equine herpesvirus type 1 in a novel hamster model. Intervirology. 2000; 43(1):27-35; ISSN: 0300-5526. Abstract.

8. van Der Meulen, K. M; Nauwynck, H. J; Buddaert, W., and Pensaert, M. B. Replication of equine herpesvirus type 1 in freshly isolated equine peripheral blood mononuclear cells and changes in susceptibility following mitogen stimulation. Journal of General Virology. 2000; 81(Pt 1):21-25; ISSN: 0022-1317. Full text.

9. van Maanen, C.; de Boer Luijtze, E., and Terpstra, C. Development and validation of a monoclonal antibody blocking ELISA for the detection of antibodies against both equine herpesvirus type 1 (EHV1) and equine herpesvirus type 4 (EHV4). Veterinary Microbiology. 2000; 71( 1-2):37-51; ISSN: 0378-1135. Abstract.

10. Walker, C.; Ruitenberg, K. M; Love, D. N, and Millar Whalley, J. Immunization of BALB/c mice with DNA encoding equine herpesvirus 1 (EHV-1) glycoprotein D affords partial protection in a model of EHV-1-induced abortion. Veterinary Microbiology. 2000; 76(3):211-220; ISSN: 0378-1135. Abstract.

11. Yasunaga, S. ; Maeda, K.; Matsumura, T.; Kondo, T., and Kai, K. Application of a type-specific enzyme-linked immunosorbent assay for equine herpesvirus types 1 and 4 (EHV-1 and -4) to horse populations inoculated with inactivated EHV-1 vaccine. Journal of Veterinary Medical Science. 2000; 62(7):687-691; ISSN: 0916-7250. Full text.

12. Zhang, Y.; Smith, P. M; Jennings, S. R, and O'Callaghan, D. J. Quantitation of virus-specific classes of antibodies following immunization of mice with attenuated equine herpesvirus 1 and viral glycoprotein D. Virology. 2000; 268(2):482-492; ISSN: 0042-6822. Abstract.

13. Breathnach, C. C; Yeargan, M. R; Sheoran, A. S, and Allen, G. P. The mucosal humoral immune response of the horse to infective challenge and vaccination with equine herpesvirus-1 antigens. Equine Veterinary Journal. 2001; 33(7):651-657; ISSN: 0425-1644. Abstract.

14. Heldens, J. G ; Hannant, D.; Cullinane, A. A; Prendergast, M. J; Mumford, J. A; Nelly, M.; Kydd, J. H; Weststrate, M. W, and van den Hoven, R. Clinical and virological evaluation of the efficacy of an inactivated EHV1 and EHV4 whole virus vaccine (Duvaxyn EHV1,4). Vaccination/challenge experiments in foals and pregnant mares. Vaccine. 2001; 19(30):4307-4317; ISSN: 0264-410X. Abstract.

15. Heldens, J. G; Kersten, A. J; Weststrate, M. W, and van den Hoven, R. Duration of immunity induced by an adjuvanted and inactivated equine influenza, tetanus and equine herpesvirus 1 and 4 combination vaccine. Veterinary Quarterly. 2001; 23(4):210-217; ISSN: 0165-2176. Abstract.

16. Osterrieder, N.; Seyboldt, C., and Elbers, K. Deletion of gene 52 encoding glycoprotein M of equine herpesvirus type 1 strain RacH results in increased immunogenicity. Veterinary Microbiology. 2001; 81(3):219-226; ISSN: 0378-1135. Abstract.

17. Ruitenberg, K. M; Gilkerson, J. R; Wellington, J. E; Love, D. N, and Whalley, J. M. Equine herpesvirus 1 glycoprotein D expressed in Pichia pastoris is hyperglycosylated and elicits a protective immune response in the mouse model of EHV-1 disease. Virus Research. 2001; 79( 1-2):125-135; ISSN: 0168-1702. Abstract.

18. Singh, B. K; Yadav, M. P, and Tewari, S. C. Neutralizing and complement-fixing monoclonal antibodies as an aid to the diagnosis of equine herpesvirus-1 infection. Veterinary Research Communications. 2001; 25(8):675-686; ISSN: 0165-7380. Abstract.

19. van der Meulen, K. M; Nauwynck, H. J, and Pensaert, M. B. Mitogen stimulation favours replication of equine herpesvirus-1 in equine blood mononuclear cells by inducing cell proliferation and formation of close intercellular contacts. Journal of General Virology. 2001; 82(Pt 8):1951-1957; ISSN: 0022-1317. Full text.

20. Allen, G.P., “Epidemic disease caused by Equine herpesvirus-1: recommendations for prevention and control,” Equine Veterinary Education, 2002, Vol 4, p 177-184. (pdf file).

21. Foote, C. E; Love, D. N; Gilkerson, J. R, and Whalley, J. M . Serological responses of mares and weanlings following vaccination with an inactivated whole virus equine herpesvirus 1 and equine herpesvirus 4 vaccine. Veterinary Microbiology. 2002; 88(1):13-25; ISSN: 0378-1135. Abstract.

22. Learmonth, G. S; Love, D. N; Wellington, J. E; Gilkerson, J. R, and Whalley, J. M. The C-terminal regions of the envelope glycoprotein gp2 of equine herpesviruses 1 and 4 are antigenically distinct. Archives of Virology. 2002; 147(3):607-615; ISSN: 0304-8608. Abstract.

23. Rudolph, J.; O'Callaghan, D. J, and Osterrieder, N. Cloning of the genomes of equine herpesvirus type 1 (EHV-1) strains KyA and racL11 as bacterial artificial chromosomes (BAC). Journal of Veterinary Medicine. B, Infectious Diseases and Veterinary Public Health. 2002 ; 49(1):31-36; ISSN: 0931-1793. Abstract.

24. Studdert, M. J. Vaccination of foals and pregnant mares with Duvaxyn EHV1, 4 vaccine. Vaccine . 2002; 20(7-8):992; ISSN: 0264-410X.

25. Taouji, S.; Collobert, C.; Gicquel, B.; Sailleau, C.; Brisseau, N.; Moussu, C.; Breuil, M. F; Pronost, S.; Borchers, K., and Zientara, S. Detection and isolation of equine herpesviruses 1 and 4 from horses in Normandy: an autopsy study of tissue distribution in relation to vaccination status. Journal of Veterinary Medicine. B, Infectious Diseases and Veterinary Public Health. 2002; 49(8):394-399; ISSN: 0931-1793. Abstract.

26. Foote, C. E; Gilkerson, J. R; Whalley, J. M, and Love, D. N . Seroprevalence of equine herpesvirus 1 in mares and foals on a large Hunter Valley stud farm in years pre- and postvaccination. Australian Veterinary Journal. 2003; 81(5):283-288; ISSN: 0005-0423. Full text.

27. Kydd, J. H; Wattrang, E., and Hannant, D. Pre-infection frequencies of equine herpesvirus-1 specific, cytotoxic T lymphocytes correlate with protection against abortion following experimental infection of pregnant mares. Veterinary Immunology and Immunopathology. 2003; 96(3-4):207-217; ISSN: 0165-2427. Abstract.

28. Mumford, E. L; Traub Dargatz, J. L; Carman, J.; Callan, R. J; Collins, J. K; Goltz, K. L; Romm, S. R; Tarr, S. F, and Salman, M. D. Occurrence of infectious upper respiratory tract disease and response to vaccination in horses on six sentinel premises in northern Colorado. Equine Veterinary Journal. 2003; 35(1):72-77; ISSN: 0425-1644. Abstract.

29. Patel, J. R; Bateman, H.; Williams, J., and Didlick, S. Derivation and characterisation of a live equid herpes virus-1 (EHV-1) vaccine to protect against abortion and respiratory disease due to EHV-1. Veterinary Microbiology. 2003; 91(1):23-39; ISSN: 0378-1135. Abstract.

30. Patel, J. R; Foldi, J.; Bateman, H.; Williams, J.; Didlick, S., and Stark, R. Equid herpesvirus (EHV-1) live vaccine strain C147: efficacy against respiratory diseases following EHV types 1 and 4 challenges. Veterinary Microbiology. 2003; 92(1-2):1-17; ISSN: 0378-1135. Abstract.

31. Soboll, G.; Whalley, J. M; Koen, M. T; Allen, G. P; Fraser, D. G; Macklin, M. D; Swain, W. F, and Lunn, D. P. Identification of equine herpesvirus-1 antigens recognized by cytotoxic T lymphocytes. Journal of General Virology. 2003; 84(Pt 10):2625-2634; ISSN: 0022-1317. Abstract.

32. Szeredi, L.; Aupperle, H., and Steiger, K. Detection of equine herpesvirus-1 in the fetal membranes of aborted equine fetuses by immunohistochemical and in-situ hybridization techniques. Journal of Comparative Pathology. 2003; 129(2-3):147-153; ISSN: 0021-9975. Abstract.

33. van der Meulen, K. M; Nauwynck, H. J, and Pensaert, M. B. Absence of viral antigens on the surface of equine herpesvirus-1-infected peripheral blood mononuclear cells: a strategy to avoid complement-mediated lysis. Journal of General Virology. 2003; 84(Pt 1):93-97; ISSN: 0022-1317. Full text.

34. Zhang, Y.; Smith, P. M; Frampton, A. R; Osterrieder, N.; Jennings, S. R, and O'Callaghan, D. J. Cytokine profiles and long-term virus-specific antibodies following immunization of CBA mice with equine herpesvirus 1 and viral glycoprotein D. Viral Immunology. 2003; 16(3):307-320; ISSN: 0882-8245. Abstract.

35. Foote, C. E; Love, D. N; Gilkerson, J. R, and Whalley, J. M . Detection of EHV-1 and EHV-4 DNA in unweaned Thoroughbred foals from vaccinated mares on a large stud farm. Equine Veterinary Journal. 2004; 36(4):341-345; ISSN: 0425-1644. Abstract.

36. Kondo, T.; McGregor, M.; Chu, Q.; Chen, D.; Horimoto, T., and Kawaoka, Y. A protective effect of epidermal powder immunization in a mouse model of equine herpesvirus-1 infection. Virology. 2004; 318(1):414-419; ISSN: 0042-6822. Abstract.

37. Maeda, K.; Kai, K.; Hayashi, T.; Hasegawa, K., and Matsumura, T. Intercellular adhesion molecule-1 (ICAM-1) and lymphocyte function-associated antigen-1 (LFA-1) contribute to the elimination of equine herpesvirus type 1 (EHV-1) from the lungs of intranasally infected BALB/c mice. Journal of Comparative Pathology. 2004; 130(2-3):162-170; ISSN: 0021-9975. Abstract.

38. Minke, J. M; Audonnet, J. C, and Fischer, L. Equine viral vaccines: the past, present and future. Veterinary Research. 2004; 35(4):425-443; ISSN: 0928-4249. Full text.

39. Newton, J. R ; Geraghty, R. J; Castillo Olivares, J.; Cardwell, J. M, and Mumford, J. A. Evidence that use of an inactivated equine herpesvirus vaccine induces serum cytotoxicity affecting the equine arteritis virus neutralisation test. Vaccine. 2004; 22(29-30):4117-4123; ISSN: 0264-410X. Abstract.

40. Patel, J. R; Didlick, S., and Bateman, H. Efficacy of a live equine herpesvirus-1 (EHV-1) strain C147 vaccine in foals with maternally-derived antibody: protection against EHV-1 infection. Equine Veterinary Journal. 2004; 36(5):447-451; ISSN: 0425-1644. Abstract.

41. Reed, S. M and Toribio, R. E. Equine herpesvirus 1 and 4. Veterinary Clinics of North America. Equine Practice. 2004; 20(3):631-642; ISSN: 0749-0739. Abstract.

42. Singh, B. K; Ahuja, S., and Gulati, B. R. Development of a neutralizing monoclonal antibody-based blocking ELISA for detection of equine herpesvirus 1 antibodies. Veterinary Research Communications. 2004; 28(5):437-446; ISSN: 0165-7380. Abstract.

43. Breathnach, C. C.; Soboll, G.; Suresh, M., and Lunn, D. P. Equine herpesvirus-1 infection induces IFN-gamma production by equine T lymphocyte subsets. Veterinary Immunology and Immunopathology. 2005; 103(3-4):207-215; ISSN: 0165-2427. Abstract.  

44. Foote, C. E; Love, D. N; Gilkerson, J. R; Rota, J.; Trevor Jones, P.; Ruitenberg, K. M; Wellington, J. E, and Whalley, J. M. Serum antibody responses to equine herpesvirus 1 glycoprotein D in horses, pregnant mares and young foals. Veterinary Immunology and Immunopathology. 2005; 105(1-2):47-57; ISSN: 0165-2427. Abstract.

45. Paillot, R.; Daly, J. M; Juillard, V.; Minke, J. M; Hannant, D., and Kydd, J. H. Equine interferon gamma synthesis in lymphocytes after in vivo infection and in vitro stimulation with EHV-1. Vaccine. 2005; 23(36):4541-4551; ISSN: 0264-410X. Abstract.

46. Patel, J. R and Heldens, J. Equine herpesviruses 1 (EHV-1) and 4 (EHV-4)--epidemiology, disease and immunoprophylaxis: a brief review. Veterinary Journal. 2005; 170(1):14-23; ISSN: 1090-0233. Abstract.

47. Trapp, S.; von Einem, J.; Hofmann, H.; Kostler, J.; Wild, J.; Wagner, R.; Beer, M., and Osterrieder, N. Potential of equine herpesvirus 1 as a vector for immunization. Journal of Virology. 2005; 79(9):5445-5454; ISSN: 0022-538X. Full text.

48. Coombs, D. K.; Patton, T.; Kohler, A. K.; Soboll, G.; Breathnach, C.; Townsend, H. G., and Lunn, D. P. Cytokine responses to EHV-1 infection in immune and non-immune ponies. Veterinary Immunology and Immunopathology. 2006; 111(1-2):109-116; ISSN: 0165-2427. Abstract.

49. Foote, C. E.; Love, D. N.; Gilkerson, J. R.; Wellington, J. E., and Whalley, J. M. EHV-1 and EHV-4 infection in vaccinated mares and their foals. Veterinary Immunology and Immunopathology. 2006; 111(1-2):41-46; ISSN: 0165-2427. Abstract.

50. Goodman, L. B.; Wagner, B.; Flaminio, M. J.; Sussman, K. H.; Metzger, S. M.; Holland, R., and Osterrieder, N. Comparison of the efficacy of inactivated combination and modified-live virus vaccines against challenge infection with neuropathogenic equine herpesvirus type 1 (EHV-1). Vaccine. 2006; 24(17):3636-3645; ISSN: 0264-410X. Abstract.

51. Holmes, M. A.; Townsend, H. G.; Kohler, A. K.; Hussey, S.; Breathnach, C.; Barnett, C.; Holland, R., and Lunn, D. P. Immune responses to commercial equine vaccines against equine herpesvirus-1, equine influenza virus, eastern equine encephalomyelitis, and tetanus. Veterinary Immunology and Immunopathology. 2006; 111(1-2):67-80; ISSN: 0165-2427. Abstract.

52. Kydd, J. H.; Townsend, H. G., and Hannant, D. The equine immune response to equine herpesvirus-1: the virus and its vaccines. Veterinary Immunology and Immunopathology. 2006; 111(1-2):15-30; ISSN: 0165-2427. Abstract.

53. Minke, J. M.; Fischer, L.; Baudu, P.; Guigal, P. M.; Sindle, T.; Mumford, J. A., and Audonnet, J. C. Use of DNA and recombinant canarypox viral (ALVAC) vectors for equine herpes virus vaccination. Veterinary Immunology and Immunopathology. 2006; 111(1-2):47-57; ISSN: 0165-2427. Abstract.

54. Paillot, R.; Ellis, S. A.; Daly, J. M.; Audonnet, J. C.; Minke, J. M.; Davis Poynter, N.; Hannant, D., and Kydd, J. H. Characterisation of CTL and IFN-gamma synthesis in ponies following vaccination with a NYVAC-based construct coding for EHV-1 immediate early gene, followed by challenge infection. Vaccine. 2006; 24(10):1490-1500; ISSN: 0264-410X. Abstract.

55. Rosas, C. T.; Goodman, L. B.; von Einem, J., and Osterrieder, N. Equine herpesvirus type 1 modified live virus vaccines: quo vaditis? Expert Review of Vaccines. 2006; 5(1):119-131. Abstract.

56. Soboll, G.; Hussey, S. B.; Whalley, J. M.; Allen, G. P.; Koen, M. T.; Santucci, N.; Fraser, D. G.; Macklin, M. D.; Swain, W. F., and Lunn, D. P. Antibody and cellular immune responses following DNA vaccination and EHV-1 infection of ponies. Veterinary Immunology and Immunopathology. 2006; 111(1-2):81-95; ISSN: 0165-2427. Abstract.

57. Tsujimura, K.; Yamanaka, T.; Kondo, T.; Fukushi, H., and Matsumura, T. Pathogenicity and immunogenicity of equine herpesvirus type 1 mutants defective in either gI or gE gene in murine and hamster models. Journal of Veterinary Medical Science. 2006; 68(10):1029-1038; ISSN: 0916-7250. Full text (pdf).

58. Weerasinghe, C. U.; Learmonth, G. S.; Gilkerson, J. R.; Foote, C. E.; Wellington, J. E. , and Whalley, J. M. Equine herpesvirus 1 glycoprotein D expressed in E. coli provides partial protection against equine herpesvirus infection in mice and elicits virus-neutralizing antibodies in the horse. Veterinary Immunology and Immunopathology. 2006; 111(1-2):59-66; ISSN: 0165-2427. Abstract.

59. Rosas, C. T.; Konig, P.; Beer, M.; Dubovi, E. J.; Tischer, B. K, and Osterrieder, N. Evaluation of the vaccine potential of an equine herpesvirus type 1 vector expressing bovine viral diarrhea virus structural proteins.  Journal of General Virology. 2007; 88(3):748-757; ISSN: 0022-1317. Abstract.

top of bibliographies

I. Therapy and Management

1. Frampton, A. R Jr; Smith, P. M; Zhang, Y.; Matsumura, T.; Osterrieder, N., and O'Callaghan, D. J. Contribution of gene products encoded within the unique short segment of equine herpesvirus 1 to virulence in a murine model. Virus Research. 2002; 90(1-2):287-301; ISSN: 0168-1702. Abstract.

2. Bentz, B. G.; Maxwell, L. K.; Erkert, R. S.; Royer, C. M.; Davis, M. S.; MacAllister, C. G., and Clarke, C. R. Pharmacokinetics of acyclovir after single intravenous and oral administration to adult horses. Journal of Veterinary Internal Medicine. 2006; 20(3):589-594; ISSN: 0891-6640. Abstract.

3. Field, H. J.; Biswas, S., and Mohammad, I. T. Herpesvirus latency and therapy--from a veterinary perspective. Antiviral Research. 2006; 71(2-3):127-133; ISSN: 0166-3542. Abstract.

4. Foote, C. E.; Raidal, S. L.; Pecenpetelovska, G.; Wellington, J. E., and Whalley, J. M. Inoculation of mares and very young foals with EHV-1 glycoproteins D and B reduces virus shedding following respiratory challenge with EHV-1. Veterinary Immunology and Immunopathology. 2006; 111(1-2):97-108; ISSN: 0165-2427. Abstract.

5. Henninger, R. W.; Reed, S. M.; Saville, W. J.; Allen, G. P.; Hass, G. F.; Kohn, C. W., and Sofaly, C. Outbreak of neurologic disease caused by equine herpesvirus-1 at a university equestrian center. Journal of Veterinary Internal Medicine. 2007; 21(1):157-165; ISSN: 0891-6640. Abstract.

6. Irwin, V. L.; Traub Dargatz, J. L.; Newton, J. R.; Scase, T. J.; Davis Poynter, N. J.; Nugent, J.; Creis, L.; Leaman, T. R., and Smith, K. C. Investigation and management of an outbreak of abortion related to equine herpesvirus type 1 in unvaccinated ponies. Veterinary Record. 2007; 160(11):378-380; ISSN: 0042-4900.

top of bibliographies

J. Herpes Virology

1. Bowles, D. E; Kim, S. K, and O'Callaghan, D. J. Characterization of the trans-activation properties of equine herpesvirus 1 EICP0 protein. Journal of Virology. 2000; 74(3):1200-1208; ISSN: 0022-538X. Full text.

2. Carvalho, R.; Oliveira, A. M; Souza, A. M; Passos, L. M, and Martins, A. S. Prevalence of equine herpesvirus type 1 latency detected by polymerase chain reaction. Archives of Virology. 2000; 145(9):1773-1787; ISSN: 0304-8608. Abstract.

3. Carvalho, R.; Passos, L. M, and Martins, A. S. Development of a differential multiplex PCR assay for equine herpesvirus 1 and 4 as a diagnostic tool. Journal of Veterinary Medicine. B, Infectious Diseases and Veterinary Public Health. 2000 ; 47(5):351-359; ISSN: 0931-1793. Abstract.

4. Chesters, P. M; Hughes, A., and Edington, N. Equid herpesvirus 1: platelets and alveolar macrophages are potential sources of activated TGF-B1 in the horse. Veterinary Immunology and Immunopathology. 2000; 75(1-2):71-79; ISSN: 0165-2427. Abstract.

5. Derbigny, W. A; Kim, S. K; Caughman, G. B, and O'Callaghan, D. J. The EICP22 protein of equine herpesvirus 1 physically interacts with the immediate-early protein and with itself to form dimers and higher-order complexes. Journal of Virology. 2000; 74(3):1425-1435; ISSN: 0022-538X. Full text.

6. Grapes, M. and O'Hare, P. Differences in determinants required for complex formation and transactivation in related VP16 proteins. Journal of Virology. 2000; 74(21):10112-10121; ISSN: 0022-538X. Full text.

7. Koen, M. T; Walker, C.; Wellington, J. E; Love, D. N, and Whalley, J. M. Characterisation of IE and UL5 gene products of equine herpesvirus 1 using DNA inoculation of mice. Archives of Virology. 2000; 145(12):2677-2686; ISSN: 0304-8608. Abstract.

8. Seyboldt, C. ; Granzow, H., and Osterrieder, N. Equine herpesvirus 1 (EHV-1) glycoprotein M: effect of deletions of transmembrane domains. Virology. 2000; 278(2):477-489; ISSN: 0042-6822. Abstract.

9. Galosi, C. M ; Vila Roza, M. V; Oliva, G. A; Pecoraro, M. R ; Echeverria, M. G; Corva, S., and Etcheverrigaray, M. E. A polymerase chain reaction for detection of equine herpesvirus-1 in routine diagnostic submissions of tissues from aborted foetuses. Journal of Veterinary Medicine. B, Infectious Diseases and Veterinary Public Health. 2001; 48(5):341-346; ISSN: 0931-1793. Abstract.

10. Iqbal, J.; Purewal, A. S, and Edington, N. EHV-1 gene63 is not essential for in vivo replication in horses and mice, nor does it affect reactivation in the horse: short communication. Acta Veterinaria Hungarica. 2001; 49(4):473-478; ISSN: 0236-6290. Abstract.

11. Jang, H. K; Albrecht, R. A; Buczynski, K. A; Kim, S. K; Derbigny, W. A, and O'Callaghan, D. J. Mapping the sequences that mediate interaction of the equine herpesvirus 1 immediate-early protein and human TFIIB. Journal of Virology. 2001; 75(21):10219-10230; ISSN: 0022-538X. Full text.

12. Kim, S. K; Buczynski, K. A; Caughman, G. B, and O'Callaghan, D. J. The equine herpesvirus 1 immediate-early protein interacts with EAP, a nucleolar-ribosomal protein. Virology. 2001; 279(1):173-184; ISSN: 0042-6822. Abstract.

13. Kim, S. K and O'Callaghan, D. J. Molecular characterizations of the equine herpesvirus 1 ETIF promoter region and translation initiation site. Virology. 2001; 286(1):237-247; ISSN: 0042-6822. Abstract.

14. Oettler, D.; Kaaden, O. R, and Neubauer, A. The equine herpesvirus 1 UL45 homolog encodes a glycosylated type II transmembrane protein and is involved in virus egress. Virology. 2001; 279(1):302-312; ISSN: 0042-6822. Abstract.

15. Osterrieder, N.; Seyboldt, C., and Elbers, K. Deletion of gene 52 encoding glycoprotein M of equine herpesvirus type 1 strain RacH results in increased immunogenicity. Veterinary Microbiology. 2001; 81(3):219-226; ISSN: 0378-1135. Abstract.

16. Ruitenberg, K. M; Gilkerson, J. R; Wellington, J. E; Love, D. N, and Whalley, J. M. Equine herpesvirus 1 glycoprotein D expressed in Pichia pastoris is hyperglycosylated and elicits a protective immune response in the mouse model of EHV-1 disease. Virus Research. 2001; 79( 1-2):125-135; ISSN: 0168-1702. Abstract.

17. Smith, D.; Hamblin, A., and Edington, N. Infection of endothelial cells with equine herpesvirus-1 (EHV-1) occurs where there is activation of putative adhesion molecules: a mechanism for transfer of virus. Equine Veterinary Journal. 2001; 33(2):138-142; ISSN: 0425-1644. Abstract.

18. van der Meulen, K. M; Nauwynck, H. J, and Pensaert, M. B. Mitogen stimulation favours replication of equine herpesvirus-1 in equine blood mononuclear cells by inducing cell proliferation and formation of close intercellular contacts. Journal of General Virology. 2001; 82(Pt 8):1951-1957; ISSN: 0022-1317. Full text.

19. Derbigny, W. A; Kim, S. K; Jang, H. K, and O'Callaghan, D. J. EHV-1 EICP22 protein sequences that mediate its physical interaction with the immediate-early protein are not sufficient to enhance the trans-activation activity of the IE protein. Virus Research. 2002; 84(1-2):1-15; ISSN: 0168-1702. Abstract.

20. Huang Ja, J. A; Ficorilli, N.; Hartley, C. A; Allen, G. P, and Studdert, M. J. Polymorphism of open reading frame 71 of equine herpesvirus-4 (EHV-4) and EHV-1. Journal of General Virology. 2002; 83( Pt 3):525-531; ISSN: 0022-1317. Full text.

21. Iqbal, J. and Edington, N. Equid herpesvirus 1 is neurotropic in mice, but latency from which infectious virus can be reactivated does not occur. Acta Veterinaria Hungarica. 2002; 50(1):117-129; ISSN: 0236-6290. Abstract.

22. Neubauer, A. ; Rudolph, J.; Brandmuller, C.; Just, F. T, and Osterrieder, N. The equine herpesvirus 1 UL34 gene product is involved in an early step in virus egress and can be efficiently replaced by a UL34-GFP fusion protein. Virology. 2002; 300(2):189-204; ISSN: 0042-6822. Abstract.

23. Rudolph, J.; O'Callaghan, D. J, and Osterrieder, N. Cloning of the genomes of equine herpesvirus type 1 (EHV-1) strains KyA and racL11 as bacterial artificial chromosomes (BAC). Journal of Veterinary Medicine. B, Infectious Diseases and Veterinary Public Health. 2002 ; 49(1):31-36; ISSN: 0931-1793. Abstract.

24. Rudolph, J. and Osterrieder, N. Equine herpesvirus type 1 devoid of gM and gp2 is severely impaired in virus egress but not direct cell-to-cell spread. Virology. 2002; 293(2):356-367; ISSN: 0042-6822. Abstract.

25. Rudolph, J.; Seyboldt, C.; Granzow, H., and Osterrieder, N. The gene 10 (UL49.5) product of equine herpesvirus 1 is necessary and sufficient for functional processing of glycoprotein M. Journal of Virology. 2002; 76(6):2952-2963; ISSN: 0022-538X. Full text.

26. Smith, D.; Hamblin, A., and Edington, N. Equid herpesvirus 1 infection of endothelial cells requires activation of putative adhesion molecules: an in vitro model. Clinical and Experimental Immunology. 2002; 129(2):281-287; ISSN: 0009-9104. Abstract.

27. van der Meulen, K. M; Nauwynck, H. J, and Pensaert, M. B. Increased susceptibility of peripheral blood mononuclear cells to equine herpes virus type 1 infection upon mitogen stimulation: a role of the cell cycle and of cell-to-cell transmission of the virus. Veterinary Microbiology. 2002; 86(1-2):157-163; ISSN: 0378-1135. Abstract.

28. Bryant, N. A; Davis Poynter, N.; Vanderplasschen, A., and Alcami, A. Glycoprotein G isoforms from some alphaherpesviruses function as broad-spectrum chemokine binding proteins. EMBO Journal. 2003; 22(4):833-846; ISSN: 0261-4189. Abstract.

29. Kim, S. K; Jang, H. K; Albrecht, R. A; Derbigny, W. A; Zhang, Y., and O'Callaghan, D. J. Interaction of the equine herpesvirus 1 EICP0 protein with the immediate-early (IE) protein, TFIIB, and TBP may mediate the antagonism between the IE and EICP0 proteins. Journal of Virology. 2003; 77(4):2675-2685; ISSN: 0022-538X. Full text.

30. Kirisawa, R. ; Kobayashi, T.; Uematsu, R.; Ikeda, A.; Kuroiwa, R.; Urakami, A., and Iwai, H. Growth of recombinant equine herpesvirus 1 (EHV-1) replaced with passage-induced mutant gene 1 and gene 71 derived from an attenuated EHV-1 in cell cultures and in the lungs of mice. Veterinary Microbiology. 2003; 95(3):159-174; ISSN: 0378-1135. Abstract.

31. Patel, J. R; Bateman, H.; Williams, J., and Didlick, S. Derivation and characterisation of a live equid herpes virus-1 (EHV-1) vaccine to protect against abortion and respiratory disease due to EHV-1. Veterinary Microbiology. 2003; 91(1):23-39; ISSN: 0378-1135. Abstract.

32. Schimmer, C. and Neubauer, A. The equine herpesvirus 1 UL11 gene product localizes to the trans-golgi network and is involved in cell-to-cell spread. Virology. 2003; 308(1):23-36; ISSN: 0042-6822. Abstract.

33. Szeredi, L.; Palfi, V., and Molnar, T. Comparison of methods for the diagnosis of equine herpesvirus type 1 infection. Acta Veterinaria Hungarica. 2003; 51(2):153-163; ISSN: 0236-6290. Abstract.

34. van der Meulen, K. M; Nauwynck, H. J, and Pensaert, M. B. Absence of viral antigens on the surface of equine herpesvirus-1-infected peripheral blood mononuclear cells: a strategy to avoid complement-mediated lysis. Journal of General Virology. 2003; 84(Pt 1):93-97; ISSN: 0022-1317. Full text.

35. Yao, H.; Osterrieder, N., and O'Callaghan, D. J. Generation and characterization of an EICP0 null mutant of equine herpesvirus 1. Virus Research. 2003; 98(2):163-172; ISSN: 0168-1702. Abstract.

36. Albrecht, R. A; Kim, S. K; Zhang, Y.; Zhao, Y., and O'Callaghan, D. J. The equine herpesvirus 1 EICP27 protein enhances gene expression via an interaction with TATA box-binding protein. Virology. 2004; 324(2):311-326; ISSN: 0042-6822. Abstract.

37. Ambagala, A. P; Gopinath, R. S, and Srikumaran, S. Peptide transport activity of the transporter associated with antigen processing (TAP) is inhibited by an early protein of equine herpesvirus-1. Journal of General Virology. 2004; 85(Pt 2):349-353; ISSN: 0022-1317. Full text.

38. Chvala, S.; Nowotny, N.; Kotzab, E.; Cain, M., and van den Hoven, R. Use of the meridian test for the detection of equine herpesvirus type 1 infection in horses with decreased performance. Journal of the American Veterinary Medical Association. 2004; 225(4):554-559; ISSN: 0003-1488. Abstract.

39. Ibrahim, el S. M.; Pagmajav, O.; Yamaguchi, T.; Matsumura, T., and Fukushi, H. Growth and virulence alterations of equine herpesvirus 1 by insertion of a green fluorescent protein gene in the intergenic region between ORFs 62 and 63. Microbiology and Immunology. 2004; 48(11):831-842; ISSN: 0385-5600. Full text.

40. Kim, S. K; Albrecht, R. A, and O'Callaghan, D. J. A negative regulatory element (base pairs -204 to -177) of the EICP0 promoter of equine herpesvirus 1 abolishes the EICP0 protein's trans-activation of its own promoter. Journal of Virology. 2004; 78(21):11696-11706; ISSN: 0022-538X. Full text.

41. von Einem, J.; Wellington, J.; Whalley, J. M; Osterrieder, K.; O'Callaghan, D. J, and Osterrieder, N. The truncated form of glycoprotein gp2 of equine herpesvirus 1 (EHV-1) vaccine strain KyA is not functionally equivalent to full-length gp2 encoded by EHV-1 wild-type strain RacL11. Journal of Virology. 2004; 78( 6):3003-3013; ISSN: 0022-538X. Full text.

42. Albrecht, R. A; Kim, S. K, and O'Callaghan, D. J. The EICP27 protein of equine herpesvirus 1 is recruited to viral promoters by its interaction with the immediate-early protein. Virology. 2005; 333(1):74-87; ISSN: 0042-6822. Abstract.

43. Breathnach, C. C; Soboll, G.; Suresh, M., and Lunn, D. P. Equine herpesvirus-1 infection induces IFN-gamma production by equine T lymphocyte subsets. Veterinary Immunology and Immunopathology. 2005; 103(3-4):207-215; ISSN: 0165-2427. Abstract.

44. Buczynski, K. A.; Kim, S. K., and O' Callaghan, D. J. Initial characterization of 17 viruses harboring mutant forms of the immediate-early gene of equine herpesvirus 1. Virus Genes. 2005; 31(2):229-239; ISSN: 0920-8569. Abstract.

45. Foote, C. E; Love, D. N; Gilkerson, J. R; Rota, J.; Trevor Jones, P.; Ruitenberg, K. M; Wellington, J. E, and Whalley, J. M. Serum antibody responses to equine herpesvirus 1 glycoprotein D in horses, pregnant mares and young foals. Veterinary Immunology and Immunopathology. 2005; 105(1-2):47-57; ISSN: 0165-2427. Abstract.

46. Frampton, A. R. Jr; Goins, W. F.; Cohen, J. B.; von Einem, J.; Osterrieder, N.; O'Callaghan, D. J., and Glorioso, J. C. Equine herpesvirus 1 utilizes a novel herpesvirus entry receptor. Journal of Virology. 2005; 79(5):3169-3173; ISSN: 0022-538X. Full text (pdf).

47. Gupta, A. K.; Kaur, D.; Rattan, B., and Yadav, M. P. Molecular variability in different Indian isolates of equine herpesvirus-1. Veterinary Research Communications. 2005; 29(8):721-734; ISSN: 0165-7380. Abstract.

48. Hartley, C. A; Wilks, C. R; Studdert, M. J, and Gilkerson, J. R. Comparison of antibody detection assays for the diagnosis of equine herpesvirus 1 and 4 infections in horses. American Journal of Veterinary Research. 2005; 66(5):921-928; ISSN: 0002-9645. Abstract.

49. Huang, J.; Hartley, C. A.; Ficorilli, N. P.; Crabb, B. S., and Studdert, M. J. Glycoprotein G deletion mutants of equine herpesvirus 1 (EHV1; equine abortion virus) and EHV4 (equine rhinopneumonitis virus). Archives of Virology. 2005; 150(12):2583-2592; ISSN: 0304-8608. Abstract.

50. Pagamjav, O. ; Sakata, T.; Matsumura, T.; Yamaguchi, T., and Fukushi, H. Natural recombinant between equine herpesviruses 1 and 4 in the ICP4 gene. Microbiology and Immunology. 2005; 49(2):167-179; ISSN: 0385-5600. Full text.

51. Paillot, R.; Daly, J. M; Juillard, V.; Minke, J. M; Hannant, D., and Kydd, J. H. Equine interferon gamma synthesis in lymphocytes after in vivo infection and in vitro stimulation with EHV-1. Vaccine. 2005; 23(36):4541-4551; ISSN: 0264-410X. Abstract.

52. Smith, K. Equine herpesviruses 1 and 4: creeping to a solution. Veterinary Journal. 2005; 170(1):6-7; ISSN: 1090-0233.

53. Borchers, K.; Bottner, D.; Lieckfeldt, D.; Ludwig, A.; Frolich, K.; Klingeborn, B.; Widen, F.; Allen, G., and Ludwig, H. Characterization of equid herpesvirus 1 (EHV-1) related viruses from captive Grevy's zebra and blackbuck. Journal of Veterinary Medical Science. 2006; 68(7):757-760; ISSN: 0916-7250. Full text (pdf).

54. Chiam, R.; Smid, L.; Kydd, J. H.; Smith, K. C.; Platt, A., and  Davis Poynter, N. J. Use of polarised equine endothelial cell cultures and an in vitro thrombosis model for potential characterisation of EHV-1 strain variation. Veterinary Microbiology. 2006; 113(3-4):243-249; ISSN: 0378-1135. Abstract.

55. Ebner, P. D. and O'Callaghan, D. J. Genetic complexity of EHV-1 defective interfering particles and identification of novel IR4/UL5 hybrid proteins produced during persistent infection. Virus Genes. 2006; 32(3):313-320; ISSN: 0920-8569. Abstract.

56. Field, H. J.; Biswas, S., and Mohammad, I. T. Herpesvirus latency and therapy--from a veterinary perspective. Antiviral Research. 2006; 71(2-3):127-133; ISSN: 0166-3542. Abstract.

57. Galosi, C. M.; Barbeito, C. G.; Martin Ocampos, G. P.; Martinez, J. P.; Ayala, M. A.; Corva, S. G.; Fuentealba, N. A, and Gimeno, E. J. An Argentine equine herpesvirus strain with special restriction patterns protect mice challenged with a pathogenic strain. Journal of Veterinary Medicine. B. Infectious Diseases and Veterinary Public Health. 2006; 53(9):412-417; ISSN: 0931-1793. Abstract.

58. Guggemoos, S.; Just, F. T., and Neubauer, A. The equine herpesvirus 1 UL20 product interacts with glycoprotein K and promotes egress of mature particles. Journal of Virology. 2006; 80(1):95-107; ISSN: 0022-538X. Full text.

59. Hasebe, R.; Kimura, T.; Nakamura, K.; Ochiai, K.; Okazaki, K.; Wada, R., and Umemura, T. Differential susceptibility of equine and mouse brain microvascular endothelial cells to equine herpesvirus 1 infection. Archives of Virology. 2006; 151(4):775-786; ISSN: 0304-8608. Abstract.

60. Hussey, S. B.; Clark, R.; Lunn, K. F.; Breathnach, C.; Soboll, G.; Whalley, J. M., and Lunn, D. P. Detection and quantification of equine herpesvirus-1 viremia and nasal shedding by real-time polymerase chain reaction. Journal of Veterinary Diagnostic Investigation. 2006; 18(4):335-342; ISSN: 1040-6387. Abstract.

61. Kydd, J. H.; Davis Poynter, N. J.; Birch, J.; Hannant, D.; Minke, J.; Audonnet, J. C.; Antczak, D. F., and Ellis, S. A. A molecular approach to the identification of cytotoxic T-lymphocyte epitopes within equine herpesvirus 1. Journal of General Virology. 2006; 87(9):2507-2515; ISSN: 0022-1317. Abstract.

62. Loregian, A.; Case, A.; Cancellotti, E.; Valente, C.; Marsden, H. S., and Palu, G. Cloning, expression, and functional characterization of the equine herpesvirus 1 DNA polymerase and its accessory subunit. Journal of Virology. 2006; 80(13):6247-6258; ISSN: 0022-538X. Full text.

63. Reilly, F. K. Questions applications and conclusions in equine herpesvirus study. American Journal of Veterinary Research. 2006; 67(11):1820; ISSN: 0002-9645.

64. Tsujimura, K.; Yamanaka, T.; Kondo, T.; Fukushi, H., and Matsumura, T. Pathogenicity and immunogenicity of equine herpesvirus type 1 mutants defective in either gI or gE gene in murine and hamster models. Journal of Veterinary Medical Science. 2006; 68(10):1029-1038; ISSN: 0916-7250. Full text (pdf).

65. van der Meulen, K.; Caij, B.; Pensaert, M., and Nauwynck, H. Absence of viral envelope proteins in equine herpesvirus 1-infected blood nomonuclear cells during cell-associated viremia. Veterinary Microbiology. 2006; 113(3-4):265-273; ISSN: 0378-1135. Abstract.

66. van der Meulen, K. M.; Favoreel, H. W.; Pensaert, M. B., and  Nauwynck, H. J. Immune escape of equine herpesvirus 1 and other herpesviruses of veterinary importance. Veterinary Immunology and Immunopathology. 2006; 111(1-2):31-40; ISSN: 0165-2427. Abstract.

67. von Einem, J.; Schumacher, D.; O'Callaghan, D. J., and Osterrieder, N. The alpha-TIF (VP16) homologue (ETIF) of equine herpesvirus 1 is essential for secondary envelopment and virus egress. Journal of Virology. 2006; 80(6):2609-2620; ISSN: 0022-538X. Full text.

68. Ibrahim, E. S; Kinoh, M.; Matsumura, T.; Kennedy, M.; Allen, G. P.; Yamaguchi, T., and Fukushi, H. Genetic relatedness and pathogenicity of equine herpesvirus 1 isolated from onager, zebra and gazelle.  Archives of Virology. 2007; 152(2):245-255; ISSN: 0304-8608. Abstract.

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