Journal of General Virology (1992), 73, 681-686. Printed in Great Britain 681
A North American hypovirulent isolate of the chestnut blight fungus with
European isolate-related dsRNA
Bradley I. Hillman,* Yanan Tian, Peter J. Bedker and Matthew P. Brown
Department of Plant Pathology, Rutgers University, New Brunswick, New Jersey 08903, U.S.A.
We have synthesized and mapped a cDNA library
representing the one major dsRNA element associated
with hypovirulence in strain NB58 of the chestnut
blight fungus, Cryphonectria (=Endothia) parasitica,
which was isolated from recovering chestnut trees in
New Jersey, U.S.A. The linear dsRNA has a size of
approximately 12- 5 kbp and is polyadenylated at the 3'
terminus of one strand. Molecular hybridization
experiments indicate that there is sequence similarity
between the NB58 dsRNA and dsRNAs from Euro-
pean isolates of C. parasitica, but not among dsRNAs
of NB58 and those associated with other North
American isolates. Hybridization experiments with
mapped cDNA clones representing different regions of
the 12.5 kbp dsRNA indicate that the termini and the
3'-proximal two-thirds (relative to the plus strand) are
more conserved among NB58 and the European
isolates than the rest of the 5'-proximal one-third.
Nucleotide sequence analysis of the termini of NB58
dsRNA suggests common organizational features
between it and the dsRNA from French-derived strain
EP713.
Introduction
The chestnut blight fungus, Cryphonectria (=Endothia)
parasitica, was responsible for considerable damage to
European chestnut forests and virtually eliminated the
American chestnut in the first half of this century (for
reviews see Van Alfen, 1982; Anagnostakis, 1987). The
term hypovirulence generally refers to infections of the
fungus by cytoplasmic agents that can be transmitted via
hyphal anastomosis, often resulting in altered pheno-
types as well as reduced fungal virulence. Because of the
unrestrained nature of this pandemic and the aesthetic
and economic importance of the chestnut tree, a number
of debilitated, hypovirulent C. parasitica isolates of
European origin were released in the 1970s in various
parts of northeastern U.S.A., including New Jersey, in
attempts at biological control of the fungus by transmis-
sion of the hypovirulence-inducing agents. Unfortu-
nately, records identifying exactly which strains were
released are incomplete, and the investigators involved
are not available (S. L. Anagnostakis, personal commun-
ication). Previous studies regarding the interrelatedness
of dsRNAs from C. parasitica indicated that dsRNAs
from European strains are related to one another, and
dsRNAs from several North American strains are
related to one another, but that dsRNAs from European
strains are not closely related to those from North
American strains (L'Hostis et al., 1985 ; Paul et al., 1988).
The apparent absence of a protein coat surrounding
the dsRNA molecules and the inability of researchers to
infect a dsRNA-free strain of C. parasitica with any
purified form of a hypovirulence-inducing agent have
greatly hindered their study from a virological perspec-
tive. However, the weight of evidence clearly indicates
that many such agents are sufficient to cause hypoviru-
lence in the fungal host and are of viral origin (Day et al.,
1977; Anagnostakis & Day, 1979; Fulbright, 1984;
Hansen et al., 1985; Hiremath et al., 1986; Rae et al.,
1989; Choi et al., 1991a, b; Shapira et al., 1991a, b). In
the French-derived strain EP713, the easily isolated
dsRNA form of the genome is encapsulated within lipid
vesicles that appear to be devoid of a major structural
protein component, but demonstrate RNA-dependent
RNA polymerase activity (Dodds, 1980; Hansen et al.,
1985). Besides dsRNA, the cytoplasmic fraction contains
virus-specific ssRNA which is polyadenylated and
hybridizes to probes representing only one strand of the
dsRNA (Hiremath et al., 1986; Tartaglia et al., 1986).
This poly(A)-containing strand is referred to as the plus
strand, and utilizes two open reading frames (ORFs) to
express at least four polypeptides (Choi et al., 1991b;
Shapira et al., 1991b).
We are interested in comparing hypovirulence-asso-
ciated viruses of C. parasitica with the aim of understand-
ing the mechanisms of hypovirulence itself and differen-
tiating those from the hypovirulence-associated changes
in fungal morphology (Hillman et al., 1990). Strain
NB58, isolated from recovering chestnut trees in New
0001-0481 © 1992 SGM