© Benaki Phytopathological Institute
Sakr
4
2012). Consequently, the
Pl
gene life period
seems to be very short from the important
use of
Pl
gene on a large cultivated zone.
The development of sunflower resistant cul-
tivars that retain the characteristics required
for the oil crop is a lengthy process and the
rate of new
P. halstedii
virulence emergence
is more rapid than the rate that newly resis-
tant hosts can be bred. Host and race evolu-
tion in downy mildew is therefore an issue of
key agricultural and commercial importance.
In the last decade, advanced tools of bio-
technology have enabled discernment of
groups of
P. halstedii
on the molecular lev-
el and led to the shift from a morphological
to a phylogenetic species concept (Spring
and Thines, 2004). Indeed, early studies of
P.
halstedii
typically found low levels of genet-
ic diversity using RAPD markers (Komjati
et
al.,
2004), ISSR (Intelmann and Spring, 2002)
and ITS sequences (Spring and Zipper, 2006).
A study of 77 samples from twelve differ-
ent countries of six virulence races using 21
RAPD primers found low levels of differentia-
tion within and between samples grouped by
race and country (Roeckel-Drevet
et al.,
2003).
With molecular markers based on partial se-
quence of the nuclear ITS regions, Spring
et
al.
(2006) detected polymorphism between
profiles of races 100, 310 and 330, as well as
between groups of populations representing
races 700, 701, 703, 710 and 730. Giress
et al.
(2007) found high genetic variability between
the isolates from France and Russia using SNP
markers. Also, Delmotte
et al.
(2008) identi-
fied three genetically differentiated groups of
isolates organized around the first three races
described in France: 100, 710 and 703. Recent-
ly, Ahmed
et al
. (2012) suggested that multi-
ple introductions have aided in the establish-
ment of
P. halstedii
in France, and noted that
recombination facilitated by these introduc-
tions is driving the emergence of new and en-
demic races in response to host resistance.
As with the neutral markers these SNPs sep-
arated races 100 and 304 from races 703 and
710 (As-sadi
et al.,
2011). Interestingly, new in-
sights from molecular phylogenetics often
verify a narrow species concept that was pro-
posed by Gäumann in 1918 (Voglmayr and
Constantinescu, 2008).
Variation of aggressiveness in Plasmo-
para halstedii
Decomposing the pathogen cycle into
elementary life traits allows the precise
identification and quantification of differ-
ences among pathogen or host phenotypes,
but relating these life traits to the pathogen
fitness under field conditions or to the ep-
idemic development rate is complicated.
However, to characterize aggressiveness of
sunflower downy mildew populations, sun-
flower inbred lines showing different levels
of quantitative resistance and carrying no
Pl
gene should be used (Tourvieille de La-
brouhe
et al.,
2008). Four aggressiveness cri-
teria were established for
P. halstedii
races:
percentage infection, latent period,
sporu-
lation density and reduction of hypocotyl
length (dwarfing). Here, the four criteria are
presented in brief: infection was considered
as successful when the seedlings showed
sporulation of the pathogen on the shoot
surface. Latent period was defined as the
number of days of incubation necessary to
obtain the pathogen sporulating on 80% of
the plants. Sporulation density was defined
as the number of zoosporangia of the path-
ogen produced on a cotyledon.
Reduction
of hypocotyl length (dwarfing) correspond-
ed to the distance from the stem base to
cotyledon insertion and was measured af-
ter 13 days of infection on diseased plants
showing sporulation of the pathogen on the
shoot.
Differences in disease development (per-
centage of infection) could be due to differ-
ences in the rate of maturation of zoospor-
angia or in the capacity to penetrate healthy
host tissue (Delanoe, 1972). Latent period
corresponds to the time interval between
infection and appearance of symptoms on
infected plants. The number of spores pro-
duced by a diseased plant will determine
the quantity of inoculum that can infect
neighbouring plants. But they are also in-
dicative of the interaction between the host
plant and the parasite, since the quantity
of spores produced will depend on the ag-
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