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Journal of General Virology (1994), 75, 2559-2565. Printed in Great Britain 2559
Endosymbiotic bacteria associated with circulative transmission of potato
leafroll virus by Myzus persicae
Johannes F. J. M. van den Heuvel,* Martin Verbeek and Frank van der Wilk
DLO Research Institute for Plant Protection (IPO-DLO), P.O. Box 9060, 6700 GW Wageningen, The Netherlands
In order to understand the molecular mechanisms
underlying circulative transmission of potato leafroll
virus (PLRV) by aphids, we screened Myzus persicae
proteins as putative PLRV binding molecules using a
virus overlay assay of protein blots. In this way, we
found that purified PLRV particles exhibited affinity for
five aphid proteins. The one most readily detected has an
M r of 63K, and was identified as symbionin. This is the
predominant protein synthesized by the bacterial endo-
symbiont of the aphid and is released into the hae-
molymph. Since further studies clearly showed that
PLRV particles also bind to native symbionin, it was
envisaged that virus particles when acquired into the
haemocoel of an aphid interact with symbionin. In-
hibition of prokaryotic protein synthesis by feeding M.
persicae nymphs on an antibiotic-containing artificial
diet prior to PLRV acquisition reduced virus trans-
mission by more than 70 %. The major coat protein of
the virus was found to be degraded in the antibiotic-
treated aphids; this would obviously have resulted in an
increased exposure of viral RNA to enzymic breakdown
and concomitant loss of infectivity. For these reasons we
conclude that endosymbiotic bacteria play a crucial role
in determining the persistent nature of PLRV in the
aphid haemolymph and that symbionin is probably the
key protein in this interaction.
Introduction
Potato leafroll virus (PLRV) is a single-stranded RNA
virus that belongs to the luteovirus group (Waterhouse et
al., 1988). Members of this group infect a wide range of
mono- and dicotyledonous plants in which they replicate
almost exclusively in the phloem tissue. Luteoviruses are
transmitted by aphids in a circulative manner (Harrison,
1958; Sylvester, 1980). Briefly, this means that virus
particles are ingested along with phloem sap from
infected host plants and transcellularly transported
through the hindgut into the haemocoel. The acquired
virus particles are retained in an infective form in the
haemolymph for the lifespan of the aphid, apparently
without replication (Eskandari et al., 1979). Upon
contacting the accessory salivery glands, they may be
transported through this gland, eventually arriving in the
salivary duct from which they are excreted with the saliva
when the aphid feeds (Gildow & Gray, 1993). Luteo-
viruses display a high degree of vector specificity among
aphid species. These well-developed specificities suggest
an intimate association between a luteovirus and its
vectors in which both surface domains of the viral capsid
and sites or substances in the aphid are involved (Gildow,
1987). The role of the viral capsid in conferring aphid
transmissibility to a luteovirus has been convincingly
demonstrated (Rochow, 1970, 1982). However, aphid-
derived components suggested to be involved in virus
transmission have not yet been identified. Therefore, the
main objective of our work was to detect these
components and to determine their role in virus
transmission. To this end we developed an immunoblot-
based virus overlay assay to ascertain whether particles
of PLRV bind to proteinaceous components from
PLRV's major aphid vector, Myzus persicae.
Methods
Viruses and antibodies. PLRV-Wageningen (Van der Wilk et al.,
1989) was maintained on Physalis floridana as previously described
(Van den Heuvel & Peters, 1990). Beet western yellows virus (BWYV)
was provided by Dr V. Brault (Institut de Biologic Mol6culaire des
Plantes du CNRS, Strasbourg, France) and transferred to P.floridana
by M. persieae. Both viruses were purified from leaf material by a
modified enzyme-assisted procedure (Van den Heuvel et al., 1990).
Purified virus suspensions of blackeye cowpea mosaic virus (BICMV),
tomato spotted wilt virus (TSWV), cowpea mosaic virus (CPMV) and
beet necrotic yellowvein virus (BNYVV), together with their respective
antibodies, were generouslyprovided by colleagues at the Department
of Virology, Wageningen Agricultural University (WAU) and IPO-
DLO. Anti-idiotypic antibodies (AiAbs) were raised to PLRV-specific
MAbs (monoclonal antibodies) (Van den Heuvel et aL, 1990) by
injecting rabbits three times at biweeklyintervals with 1 mg of partially
purified MAbs derived from mouse ascitic fluids. Antibodies against
non-idiotypic sites of the MAbs were removed from the rabbit sera by
incubating with mouse anti-thyroglobulin antibodies and subsequent
centrifugation essentially following Hu & Rochow (1988).
Aphids and other vectors of plant viruses. M. persicae biotype WMp2
was reared on Brassica napus L. subsp, oleifera (oilseed rape) in a
0001 2519 © 1994 SGM