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Jones, 2011). Phylogenetic placement of bio-
logically defined PVY Z and PVY E isolates was
unknown until 2011, when whole genome se-
quences of PVY Z isolate L26 (Kerlan et al ., 2011)
and PVY E isolate Mon (Galvino-Costa et al .,
2011) were placed within phylogenetically de-
ined PVY NTN and a separate clade between
PVY N/NA-N and PVY NTN lineages, respectively.
PVY-L26 is a recombinant between “PVY O ” and
PVY N and PVY-Mon between PVY NTN and PVY-
NE11 (Hu et al ., 2009a,b; Galvino-Costa et al .,
2011; Kerlan et al ., 2011). The original PVY E
type isolate (Kerlan et al ., 1999; Singh et al .,
2008) remains unsequenced. However, when
the CP genes of three original PVY Z isolates were
sequenced, they grouped with “PVY O ” or PVY N-Wi
(Kehoe and Jones, 2011). Thus, PVY Z may fit
into both recombinant and non-recombinant
groupings.
Knowledge that (i) few phylogenetically de-
fined “PVY O ” isolates have been shown to belong
to biologically defined PVY O , (ii) many isolates
are recombinants between “PVY O ” and PVY N ,
and (iii) that PVY Z may fit within three distinct
phylogenetic groupings which include “PVY O
suggests that use of the name PVY O for both re-
sistance and phylogenetic PVY strain groupings
should be reconsidered. Use of PVY O as a resist-
ance grouping has historical precedence. Kehoe
and Jones (2011) suggested use of PVY O be re-
stricted to isolates defined biologically by their
reactions with gene Ny tbr , and that a letter not
used previously be employed instead (Q), giving
the designation PVY Q for the phylogenetic
grouping currently known as “PVY O ”, and the
designation PVY N:Q for the phylogenetic group-
ing PVY N:O . Kerlan et al . (2011) combined resist-
ance and phylogenetic groupings for PVY Z
isolate L26, designating it PVY Z/NTN . This approach
provides a useful means of avoiding confusion
caused by the different nomenclature systems;
for example, the historical PVY O and PVY Z iso-
lates sequenced by Kehoe and Jones (2011)
would then become PVY O/Q , PVY Z/Q , or PVY Z/N-Wi .
However, sequence characterization of whole
PVY genomes is still needed with these historical
PVY Z isolates.
Singh et al . (2008) suggested that newly
found PVY isolates should be described within
the context of the original strain groupings based
on host resistance, and not named using geograph-
ical, cultivar, or place-association designations.
However, within a strain group, amendment of
isolate names using additional codes to show
that the isolate differed at the molecular, sero-
logical, or phenotypic level was acceptable.
They recognized five strain groups based on
host resistance (PVY C , PVY O , PVY E , PVY Z ,
PVY N ). They also recognized seven strains based
on a combination of host resistance and whole
genome sequencing (PVY C , PVY O , PVY E , PVY Z ,
PVY N , PVY NTN , PVY N-Wi /PVY N:O ). The two recom-
binant strains (PVY NTN , PVY N-Wi /PVY N:O ) were
placed within strain group PVY N . They also sug-
gested that a more systematic investigation and
characterization of PVY at the biological and
molecular levels should eventually result in a
more meaningful strain concept. Karasev and
Gray (2013a) suggested an increase in the
number of PVY strains to nine (PVY C , PVY O ,
PVY E , PVY Z , PVY N , PVY N-Wi , PVY N:O , PVY NA-N ,
PVY-NE11). This change resulted from: (i) treat-
ing the ability to induce veinal necrosis in to-
bacco as a subordinate trait to reactions with
potato cultivar differentials; (ii) including
PVY NTN within PVY Z ; (iii) separating the former
PVY N-WI strain into PVY N-WI and PVY N:O ; and (iv)
adding new strains PVY NA:N and PVY-NE11.
Treating the ability to induce veinal necrosis in
tobacco as a subordinate trait was because the
viral genetic determinants involved in the in-
duction of veinal necrosis in tobacco were
shown to be distinct from the viral genetic deter-
minants involved in the induction of hypersen-
sitive resistance or PTNRD in potato (e.g. Hu
et al ., 2009b; Galvino-Costa et al ., 2011; Kerlan
et al ., 2011; Moury et al ., 2011; Faurez et al .,
2012; Tian and Valkonen, 2013). Combining
PVY NTN with PVY Z was based on findings that
potato differentials reacted similarly to both
(Barker et al ., 2009; Galvino-Costa et al ., 2011;
Kerlan et al ., 2011). Separation of the former
PVY N-WI strain into PVY N-WI and PVY N:O , and
the separate classification of PVY NA-N and PVY-
NE11, was because these strains all had mo-
lecular characteristics clearly distinct from
those of other PVY strains (Nie and Singh,
2003a; Lorenzen et al ., 2006, 2008). An im-
proved multiplex IC-RT-PCR assay was devel-
oped that distinguished these nine PVY strains
(Chikh-Ali et al ., 2013). However, PVY N-WI , PVY N:O ,
PVY NA-N , and PVY−NE11 all still lacked biological
typing data using potato cultivar differen-
tials. As mentioned above, Kehoe and Jones
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