Journal of General Virology (1993), 74, 819 828. Printed in Great Britain 819
Pathogenicity of a thymidine kinase-deficient mutant of equine
herpesvirus 1 in mice and specific pathogen-free foals
J. D. Slater, J. S. Gibson* and H. J. Field
Department of Veterinary Medicine, University of Cambridge, Madingley Road, Cambridge CB3 0ES, U.K.
Both intranasal (i.n.) and intracerebral (i.c.) inoculation
of mice with wild-type equine herpesvirus type 1 (wt
EHV-1) caused clinical signs and mortality. Virus could
be recovered from target organs (turbinates, lungs and
blood) for several days. By contrast, the thymidine
kinase (TK)-deficient deletion mutant PR1 produced
markedly less clinical disease following both i.n. and i.c.
inoculation, and, in particular, no mortality occurred.
PR1 did, however, establish productive infections
following either route of inoculation. High titres of virus
were recovered from target organs although virus did not
persist for as long as wt EHV-1 and no viraemia was
detected. Primary i.n. infection of mice with either wt
EHV-1 or PR1 protected against subsequent challenge
with wt EHV-1 5 weeks later. I.n. inoculation of specific
pathogen-free (EHV-free) foals with PR1 produced
results similar to those observed after infection of mice.
Clinical signs were milder than for wt EHV-1 and
pyrexia was short-lived or absent. PR1 could be
recovered from nasal mucus at high titres but it persisted
for only 5 days post-infection compared to 11 days in the
case of wt EHV-1. No viraemia was detected in foals
infected with PR1. On challenge with wt EHV-1, foals
given a primary infection with the mutant were partially
protected; but a viraemia with a TK + EHV-1 was
observed. These results demonstrate that our TK
mutant PR1 is markedly less pathogenic than wt EHV-
1, despite being able to replicate in the host. The use of
TK-deficient mutants of EHV-1 as potential vaccines in
the horse is discussed.
Introduction
The alphaherpesvirus equine herpesvirus type 1 (EHV-1)
is one of several herpesviruses known to infect horses.
EHV-1 can cause abortion and neurological disease, as
well as respiratory problems. It is a major pathogen
of horses with considerable economic and veterinary
importance (Allen & Bryans, 1986). To date, control
measures have proved inadequate, and although vaccines
are available, their efficacy is controversial (Burrows et
al., 1984; Allen & Bryans, 1986; Burki et al., 1990).
Progress towards more effective immunoprophylaxis
has been hindered by incomplete understanding of the
pathogenesis and epidemiology of EHV- 1. Our approach
to this problem has involved the use of two animal
models: colostrum-deprived specific pathogen-free
(SPF), EHV-free, foals and mice. SPF foals are taken
from their dams at birth and reared by hand using bovine
colostrum (Chong et al., 1991). They are free from EHV
infection and also from maternal antibodies. They are
invaluable in the study of EHV infection, particularly
the interaction between different EHVs (Gibson et al.,
1992a, b). Because foal production is limited and sea-
sonal, we have also developed a murine model for EHV-
1 infection whose features show remarkable similarity to
infection in the natural host (for a review, see Field et al.,
1992b).
The study of thymidine kinase (TK)-deficient mutants
of other herpesviruses has received considerable at-
tention. In particular, TK is implicated in the neuro-
pathogenicity of various alphaherpesviruses, including
herpes simplex virus types 1 and 2 (HSV-1 and HSV-2)
and suid herpesvirus type 1 (SHV-1) (Field & Wildy,
1978; Becker et al., 1984; Kit et al., 1985). Although
EHV-1 is not thought to infect neurons directly, it does
cause neurological problems in the horse, probably via
infection of endothelial cells in the central nervous
system (CNS) and subsequent ischaemic damage of
nervous tissue (Patel et al., 1982). A similar sequence of
events involving infection of placental blood vessels is
postulated in EHV-l-induced abortion.
Previous reports of EHV-1 TK-deficient mutants
suggest that they are indeed less pathogenic than the
wild-type (wt) virus (Cornick et al., 1990). The behaviour
of such mutants, however, has not been studied in detail.
Furthermore, previous work has made use of naturally
reared horses, which have the disadvantage that their
EHV status is uncertain and results obtained from them
are therefore difficult to interpret (Cornick et al., 1990).
Here we describe the characteristics of a drug-selected
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