Twelfth EuroBlight workshop

Arras (France), 3-6 May 2010

Characteristics of the Phytophthora infestans population

in Russia

N.V. Statsyuk1, M.A. Kuznetsova1, I.N. Kozlovskaya1, B.E. Kozlovsky1, S.N.

Elansky2, E.V. Morozova1, E.V. Valeva1, and A.V. Filippov1

1
All-Russian Research Institute of Phytopathology, VNIIF,
Bolshie Vyazemy, Moscow region, 143050 Russia; e-mail: natafg@rambler.ru
2
Department of Algology and Mycology, Biological Faculty,
Moscow State University, Vorob’evy Gory, Moscow, 119992 Russia

SUMMARY
A collection of 434 Phytophthora infestans isolates, obtained during 2007-2009 from potato and
tomato fields of different parts of European Russia, has been assessed for several phenotypic and
genotypic markers, including mtDNA haplotype, Pep1 and Pep2 loci, mating type, metalaxyl
sensitivity, and the virulence. The comparison of phenotypic and genotypic characteristics of P.
infestans population of the Moscow region in 2008-2009 with the similar data, obtained in 1997-
1998, has shown significant changes occurred within a 10-year period: (1) the frequency ratio of Ia
and IIa MtDNA haplotypes has shifted towards the predominance of the Ia type, (2) the percent of
strains belonged to the A2 mating type has increased, (3) the virulence gene 9 has appeared in the
current population, which now contains all 11 virulence genes, and (4) the percent of metalaxyl-
susceptible strains has significantly decreased from 78 to 58%. The analysis of the studied populations
has shown that the Leningrad, Astrakhan, and Nizhni Novgorod populations are rather uniform,
whereas the Moscow, Mariy El and Kostroma populations are characterized by a high diversity. The
Astrakhan “tomato” population has a unique pattern of virulence gene frequencies and seems to be
monoclonal.

Keywords
late blight, potato, tomato, population characteristics

INTRODUCTION
Phytophthora infestans, the causal agent of the late blight disease of potato and tomato, is the most
damaging microbial pathogen of these crops world-wide, including Russia and Eastern Europe. In
the XIX century this pathogen caused significant potato yield losses in the Northern Europe that
resulted in the Irish Potato Famine. In the last decades, new agrotechnical and plant pest control
methods, such as the use of certified seeds, breeding programs, crop rotation, and highly-effective
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fungicides, significantly decreased potato yield losses caused by the late blight. At the same time, due
to a high pathogen variability and a possible introduction of new strains by a potato shipment from

PPO-Special Report no. 14 (2010), 247- 254

247
 MATERIALS AND METHODS
central Mexico to Europe, the European population of P. infestans has undergone significant changes
P. infestans
[1-2]. The “old”isolates
populationwerewascollected
represented frombycommercial
the A1 mating potato
typeand and tomato fields, located
Ib mitochondrial in the
(Mt) DNA
following whereas
haplotype, regions of thethe European
“new” Russiaconsisted
population (Fig. 1):ofLeningrad
the isolates (21ofisolate),
both A1Moscow
and A2 mating(100 isolates,
types
including
and Ia and IIa 20 MtDNA
from tomato plants),[3-7].
haplotypes Nizhni Novgorod
Among (13 isolates),
other changes Astrakhan
in P. infestans (31 isolates
population traits,from
one
tomatoalso
should plants), Kostroma
mention (105 isolates),
an increase Smolensk
in the metalaxyl (49 isolates),
resistance and the Mariy
that decreased El Republic
the effect of the use (115
of
isolates,metalaxyl-based
popular including 93 from tomato on
fungicides plants). The
the late total control
blight number[8]. of the studied isolates was 434.
Allozyme
The interactionanalysis.
between Genotypes
strains ofatdifferent
two peptidase
mating loci types(Pep1 and Pep2)
can cause the sexualwerereproduction
analyzed using of P.a
celluloseincreasing
infestans, acetate gel theelectrophoresis
genetic diversity according to [10] this
of the progeny; withcansome modifications
possibly result in an [11]. The genetic
increase in the
diversityand
virulence for fungicide
the Pep1 locus in Russian
resistance of newly P. developed
infestans populations
strains. is significantly lower than that of
the Pep2 locus, represented by two alleles (100
In our previous studies we analyzed the phenotypic and genotypic and 112) and stained simultaneously
characteristics with the
of P. infestans
Pep1 locus (Fig. 2), so we analyzed both these markers.
populations from the Moscow region, Siberia, and Far East [9]. In this study we present the data,
The mtDNA haplotype identification was carried out according to [12] with some
obtained for the Moscow region (2008-2009) and 5 new regions, including North Western Russia
modifications.
and several regions of the Eastern part of European Russia.
The mating type was tested by the growing isolates on rye agar with the known reference
strains of the A1 and A2 mating types. Agar blocks with the studied and reference strains were
placed by pairs into Petri dishes, containing rye-vegetable agar, at a distance of 4-5 cm from
MATERIALS
each other. The AND METHODS
Petri dishes were incubated in the dark at 18С for 14 days, and then we
P.microscoped
infestans isolates
thewere
placecollected
of a hyphafrom commercial
contact betweenpotatothe andstrains
tomato to fields, located the
determine in the following
presence or
regions
absence of of
theoospores.
EuropeanIfRussia (Fig. 1):
the studied Leningrad
isolate generated(21 isolate),
oospores Moscow (100 the
only with isolates, including
A2 isolate, 20
it was
from tomato
referred plants),
to the NizhniIf Novgorod
A1 type. the isolate(13 isolates), oospores
generated Astrakhanonly (31 with
isolates
A1fromisolate,tomato
thenplants),
it was
Kostroma (105 isolates), Smolensk (49 isolates), and the Mariy El Republic
referred to the A2 type. If the isolate generated oospores with both reference strains, then (115 isolates, including
it was
93 from tomato
referred to A1A2 plants).
type. The total number of the studied isolates was 434.
Metalaxyl
Allozyme sensitivity.
analysis. Genotypes Theat two
sensitivity
peptidase of loci
isolates
(Pep1 to andmetalaxyl-containing
Pep2) were analyzed using fungicides was
a cellulose
determined
acetate by the inoculation
gel electrophoresis according of tofungicide-treated tuber discs [13]
[10] with some modifications or fungicide-containing
[11]. The genetic diversity for
medium
the [14] inwith
Pep1 locus the tested
Russian isolates
P. infestans at different
populations fungicide concentrations.
is significantly lower than that of According to the
the Pep2 locus,
obtained results,
represented by twoisolates wereand
alleles (100 considered
112) andasstained
sensitive (S), moderately
simultaneously withresistant
the Pep1(MR), locus or resistant
(Fig. 2), so
we(R).
analyzed both these markers.
The mtDNA haplotype identification was carried
out according to [12] with some modifications.
The mating type was tested by the growing isolates
on rye agar with the known reference strains of
the A1 and A2 mating types. Agar blocks with
the studied and reference strains were placed by
pairs into Petri dishes, containing rye-vegetable
agar, at a distance of 4-5 cm from each other.
The Petri dishes were incubated in the dark at
18°C or 14 days, and then we microscoped the
place of a hypha contact between the strains to
determine the presence or absence of oospores.
Fig. 2. Cellulose acetate plate showing
If the studied isolate generated oospores only
typical allozyme banding patterns of
with
two the A2 isolate,
peptidase lociit was
for referred
P. infestans to the A1
type.
isolates. Pep1 locus: 92/92 (lanes 1 and only
If the isolate generated oospores
with A1 isolate,
8); 100/100 then 5-7,
(lanes it was referred
10-12); to the A2
92/100
type.
(lanesIf the
3, isolate
4, 9). generated
Pep2 locus: oospores with both
100/100
reference
(lanes 1-5,strains,
7, 8,then it was
10-12); referred(lane
112/112 to A1A2
type.
6); 100/112 (lane 9).
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Metalaxyl sensitivity. The sensitivity of isolates

to metalaxyl-containing fungicides was
determined by the inoculation of fungicide-
Fig. 1. Locations
Fig. 1. Locations of samplingofsites.
sampling sites. treated tuber discs [13] or fungicide-containing

248
ivity of isolates to metalaxyl-containing fungicides was
fungicide-treated tuber discs [13] or fungicide-containing
es at different fungicide concentrations. According to the
dered as sensitive (S), moderately resistant (MR), or resistant
medium [14] with the tested isolates at different fungicide concentrations. According to the obtained
results, isolates were considered as sensitive (S), moderately resistant (MR), or resistant (R).

Fig. 2. Cellulose acetate plate showing typical allozyme banding patterns of two peptidase loci for P. infestans
isolates. Pep1 locus: 92/92 (lanes 1 and 8); 100/100 (lanes 5-7, 10-12); 92/100 (lanes 3, 4, 9). Pep2 locus:
Fig. 2.(lanes
100/100 Cellulose
1-5, 7, 8, acetate plate (lane
10-12); 112/112 showing
6); 100/112 (lane 9).
typical allozyme banding patterns of
Virulence. To study the virulence of isolates, we used a set of differentiator cultivars, obtained from
two peptidase loci for P. infestans
the International Potato Center (CIP, Peru) and containing 22 genotypes, including all known
isolates. Pep1 locus: 92/92 (lanes 1 and
resistance genes in different combinations. We also used the test set, containing R0-R11 genotypes
8); 100/100
and obtained from(lanes 5-7, 10-12);
the Institute of Plant92/100
Cultivation and Acclimatization (IHAR, Poland). The
(lanes 3, 4, 9). Pep2 locus:
analysis was carried out under laboratory 100/100
conditions using detached potato leaves. The leaves were
(lanes
placed into1-5,
a wet7,chamber,
8, 10-12); 112/112
representing (lane
a frame (30x40 cm) with the bottom made of a metal gause.
6);chamber
The 100/112 was(lane 9). with a glass sheet. To inoculate the leaves, we used a 4-5-day culture of
covered
P. infestans, grown on the tuber slices of a susceptible potato cultivar, or 10-12-day culture, grown
on nutrient medium. A pathogen suspension was prepared in a concentration, corresponding to
10-15 conidia in a microscopic field at 100x magnification. Tubes with the suspension (5 ml) were
sites. placed into a refrigerator (6-7ºC) to initiate the release of zoospores. The lower surface of each leaf
was inoculated by 2 drops of suspension using a micropipette; after 24 h the drops were shaken off,
and the leaves were turned upside down. The leaves were incubated at 18-20ºC. The level of the
disease development on the leaves was determined after 4-6 days of incubation. The presence of the
fruiting testified a compatible reaction and was designated by a “+”; the absence of any fruiting was
designated as “-“.

RESULTS AND DISCUSSION

Allozyme analysis.
The results of the allozyme analysis for Pep1 and Pep2 loci are shown in Fig. 3.
In all populations the predominant genotype of the Pep1 locus was 100/100; all three “tomato”
populations were represented by only this genotype. The presence of all three possible variants
(92/92, 92/100, and 100/100) was revealed only in the Moscow “potato” population, like in the case
of our previous study [9].
In the case of Pep2 locus, the genetic diversity was higher. All three possible variants were revealed in
4 populations at different proportions. Again, the most frequent genotype was 100/100, excepting the
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Leningrad population, represented by the only 112/112 genotype, Smolensk population, represented
mainly by 100/112 genotype, and Kostroma population, where the 112/112 genotype predominated.
It is interesting that the 100/100 genotype was predominant for all three tomato populations (100%
for Astrakhan and Moscow regions and 47.8% for the Mariy El Republic).

249
 100/100, excepting the Leningrad population, represented by the only 112/112 genotype,
Smolensk population, represented mainly by 100/112 genotype, and Kostroma population, where
the 112/112 genotype predominated. It is interesting that the 100/100 genotype was predominant
for all three tomato populations (100% for Astrakhan and Moscow regions and 47.8% for the
Mariy El Republic).
a b
Mariy El (t)
Mariy El (p)

Sm olensk (p)

Kostrom a (p)

Astrakhan (t)
N. Novgorod (p)

Moscow (t)

Moscow (p)
Leningrad (p)

0% 20% 40% 60% 80% 100% 0% 20% 40% 60% 80% 100%

92/92 92/100 100/100 100/100 100/112 112/112

Fig. 3. Frequency of Pep1 (a) and Pep2 (b) genotypes of P. infestans samples from different regions of European
Fig. 3. Frequency of Pep1 (a) and Pep2 (b) genotypes of P. infestans samples from different
Russia; (p) and (t) mean “potato” and “tomato” populations, respectively.
regions of European Russia; (p) and (t) mean “potato” and “tomato” populations, respectively.
Analysis of mitochondrial DNA haplotype
Analysis of mitochondrial
The resultsDNAof thehaplotype
mtDNA analysis are shown in Fig. 4.
The results of theWe
mtDNA analysis
revealed onlyaretwo
shown on Fig.haplotypes
mtDNA 4. (Ia and IIa). The Ia genotype was predominant for
We revealed only two mtDNA haplotypes (Ia and IIa). The Ia genotype was predominant for all
all populations, excepting the Mariy El “tomato” population; two populations (Leningrad and
populations, excepting the Mariy El “tomato” population; two populations (Leningrad and
Astrakhan
Astrakhan “tomato”) “tomato”)
were presented by were presented
the only by only this genotype.
this genotype.
In the case of theInMoscow
the caseregion,
of thesome
Moscow region,
changes in thesome
ratio changes in the ratio
of these MtDNA of these
haplotypes MtDNA haplotypes were
were
registered
registered comparing to thecomparing to the
previous data, previous
obtained in data, obtained
1997-1998 [9]. in 1997-1998
According to [9].
the According
earlier to the earlier study,
study, the Ia : IIa
theratio wasratio
Ia : IIa aboutwas36about
: 64, 36
whereas the recent
: 64, whereas thedata show
recent a tendency
data to the to the increase in the
show a tendency
increase in the IaIa
percentage
percentage (55(55
: 45).
: 45).

Mariy El (t)

Mariy El (p)

Kostroma (p)

Astrakhan (t)

N. Novgorod (p)

Moscow (p)

Leningrad (p)

0% 20% 40% 60% 80% 100%

Ia IIa

Fig. 4. Mitochondrial DNA haplotypes

Fig. 4. Mitochondrial P. infestans
DNAofhaplotypes of P.samples from
infestans different
samples fromregions
differentofregions of European Russia; (p)
European and
Russia; (p) and (t) mean “potato” and “tomato” populations,
(t) mean “potato” and “tomato” populations, respectively. respectively.

Mating type
Mating determined
Mating type frequencies, type for all studied populations, are shown on Table 1.
Mating
The A1 mating type type frequencies,
predominated in the most determined forpopulations;
of examined all studied populations,
in the case of are shown in Table 1.
Leningrad
The A1(tomato)
(potato) and Astrakhan mating type predominated
populations, in most
it was the of the examined
only variant revealed. Thepopulations;
A2 matingin the case of Leningrad
type prevailed in(potato)
the Nizhnii
andNovgorod,
AstrakhanSmolensk,
(tomato)and Mariy El (potato)
populations, populations
it was the (100,revealed.
only variant 94.4, The A2 mating type
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and 64.3 percents, respectively).

prevailed in thePopulations from the Moscow
Nizhnii Novgorod, Smolensk,and and
Kostroma
Mariyregions included
El (potato) a
populations (100, 94.4, and
small percent of A1A2 strains, able to generate oospores with both reference strains.
64.3 percents, respectively). Populations from the Moscow and Kostroma regions included a small
Comparing the data obtained for the Moscow region in 1997-1998 [9] and 2008-2009, one can
percent of A1A2 strains, able to generate oospores with both reference strains.
note that the A1:A2 ratio in both potato and tomato population of P. infestans changed from
Comparing
72:28 (potato) and the data
88:12 (tomato) obtained
to 61:35.4 and for the respectively,
65:35, Moscow region in percent
i.e. the 1997-1998 [9]A2
of the and 2008-2009, one can
strains increased.
250 Table 1
Occurrence of A1 and A2 mating types in Russian P. infestans populations
Region Mating type, %
A1 A2 A1A2
Leningrad (potato) 100 0 0
note that the A1:A2 ratio in both potato and tomato population of P. infestans changed from 72:28
(potato) and 88:12 (tomato) to 61:35.4 and 65:35, respectively, i.e. the percent of the A2 strains
increased.

Table 1. Occurrence of A1 and A2 mating types in Russian P. infestans populations

Mating type, %
Region
A1 A2 A1A2
Leningrad (potato) 100 0 0
Moscow (potato) 61 35.4 3.6
Moscow (tomato) 65 35 0
Nizhni Novgorod 0 100 0
Astrakhan (tomato) 100 0 0
Kostroma (potato) 80.1 19.4 0.5
Smolensk (potato) 5.6 94.4 0
Mariy El (potato) 35.7 64.3 0
Mariy El (tomato) 77.8 22.2 0

Virulence
The results of our virulence study are shown in Fig. 5.
According to our data, populations from the Kostroma and Leningrad regions were similar to each
other: the frequencies of single genes were about the same, and both populations did not include the
gene 9. Both populations have similar values of the factor of virulence (FV). The most frequent races
in both populations included 8-10 virulence genes.
Both tomato and potato populations of P. infestans from the Mariy El Republic were represented
by complex races and had the similar structure. The most frequent races were 1.2.3.4.6.7.8.10.11
(28.2%) and 1.2.3.4.5.6.7.8.10.11 (50%). The total percent of these two races in these two population
was 78.2%.
The Moscow population included all virulence genes and was presented mainly by complex races,
containing 7-11 genes (including gene 9, which was absent in the most of the studied populations). The
most frequent races were 1.3.4.7.8.10.11; 1.2.3.4.6.7.8.11; 1.2.3.4.6.7.8.10.11; 1.2.3.4.5.6.7.8.10.11;
and 1.2.3.4.5.6.7.8.9.10.11. Comparing to the earlier data, the gene 9 appeared in the population.

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251
 population was 78.2%.
The Moscow population included all virulence genes and was presented mainly by complex
races, containing 7-11 genes (including gene 9, which was absent in the most of the studied
populations). The most frequent races were 1.3.4.7.8.10.11; 1.2.3.4.6.7.8.11; 1.2.3.4.6.7.8.10.11;
1.2.3.4.5.6.7.8.10.11; and 1.2.3.4.5.6.7.8.9.10.11. Comparing to the earlier data, the gene 9
appeared in the population.

Kostrom a (p), FV = 7.2 Leningrad (p), FV = 8.2 Astrakhan (t), FV = 3.8

25 25 25

20 20 20

15 15 15

10 10 10

5 5 5

0 0 0
1 2 3 4 5 6 7 8 9 10 11 1 2 3 4 5 6 7 8 9 10 11 1 2 3 4 5 6 7 8 9 10 11

Mariy El (p), FV = 10,0 Mariy El (t), FV = 8.3

25 25

20 20

15 15

10 10

5 5

0 0
1 2 3 4 5 6 7 8 9 10 11 1 2 3 4 5 6 7 8 9 10 11

Moscow (p), FV = 8.2 Nizhnii Novgorod (p), FV = 6,3

25 25

20 20

15 15

10 10

5 5

0 0
1 2 3 4 5 6 7 8 9 10 11 1 2 3 4 5 6 7 8 9 10 11

Fig. 5. Frequencies of separate virulence genes in the studied P. infestans populations. FV, factor of virulence;
(p) and (t) mean “potato” and “tomato” populations, respectively.
The population from the Nizhnii Novgorod region consisted mainly of complex races, containing 6-11 virulence
genes (57.9%). Like the Moscow population, this population also contained the gene 9; it is also interesting
that the frequency of gene 8 was unusually low. The FV value (6.3) was lower than for the above-mentioned
populations (7.2-10.0).
In the case of the Astrakhan population, we revealed a significant predominance of virulence genes 1, 3, 4, 7.
Genes 5, 8, 10, and 11 were not revealed; the frequencies of genes 2 and 9 were very low. The FV value for
this region was very low (3.8), and the most frequent race was the race 1.3.4.7 (47%). Thus, this population
significantly differed from other ones.
In general, one should note that some of the tested populations contain the gene 9, which was not determined in
the Russian P. infestans populations during our earlier studies.

Metalaxyl resistance
The results of the analysis of the studied P. infestans populations for their metalaxyl resistance are
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shown in Fig. 6.
Both Mariy El populations were represented by only susceptible isolates; the most of isolates from
Kostroma and Moscow regions were also susceptible (98 and 58%, respectively). The percent of
susceptible strains in the Moscow “potato” population significantly decreased comparing to the

252
Metalaxyl resistance
The results of the analysis of the studied P. infestans populations for their metalaxyl resistance
are shown in Fig. 6.
Both Mariy El populations were represented by only susceptible isolates; the most of isolates
from Kostroma and Moscow regions were also susceptible (98 and 58%, respectively). The
percent earlier data [9] (from
of susceptible strains78intothe
58%). Astrakhan
Moscow “tomato”
“potato” and Leningrad
population populations
significantly decreasedwere moderately
resistant
comparing to the(100%
earlierand 76%,
data [9] respectively).
(from 78 to Finally, Nizhnii Novgorod
58%). Astrakhan “tomato” population consisted mainly of
and Leningrad
populations were moderately
the resistant isolates resistant
(73%). (100% and 76%, respectively). Finally, Nizhnii Novgorod
population consisted mainly of the resistant isolates (73%).

Mariy El (t)

Mariy El (p)

Kostroma (p)
S
Astrakhan (t) MR
N. Novgorod (p) R

Moscow (p)

Leningrad (p)

0% 20% 40% 60% 80% 100%

Fig. Fig.
6. Frequency of P.ofinfestans
6. Frequency strains
P. infestans withwith
strains different level
different of the
level metalaxyl
of the metalaxyl resistance
resistanceinin different regions of
differentEuropean
regions of European
Russia. Russia. S,
S, susceptible susceptible
strains, strains, MR,
MR, moderately moderately
resistant strains,resistant strains,
R, resistant R, (p) and (t) mean
strains;
resistant
“potato” strains; (p) andpopulations,
and “tomato” (t) mean “potato” and “tomato” populations, respectively.
respectively.
CONCLUSIONS
CONCLUSIONS
The comparison of phenotypic and genotypic characteristics of P. infestans population of the
MoscowThe comparison
region in 2008-2009of phenotypic and data,
with the similar genotypic characteristics
obtained in 1997-1998ofshowedP. infestans population of the
that some
Moscow
significant changesregion in 2008-2009
occurred withperiod.
within a 10-year the similar data, obtained
The frequency in and
ratio of Ia 1997-1998
IIa MtDNA showed that some
haplotypes shifted from
significant 36:64
changes to 55:45,within
occurred increasing the percent
a 10-year period.ofThe
Ia type. The A1:A2
frequency ratio
ratio of Ia in
and IIa MtDNA
both potato and tomato
haplotypes P. infestans
shifted populations
from 36:64 to 55:45,ofincreasing
the regiontheshifted from
percent of72:28 (potato)
Ia type. and ratio in both
The A1:A2
88:12 (tomato)
potato toand61:35.4
tomatoandP.65:35, respectively,
infestans i.e. the
populations percent
of the regionof A2 strains
shifted increased.
from 72:28 The
(potato) and 88:12
virulence gene 9 appeared in the current population, which now contains all 11 virulence genes.
(tomato) to 61:35.4 and 65:35, respectively, i.e. the percent of A2 strains increased. The virulence
The percent of susceptible strains in the “potato” population decreased from 78 to 58%.
gene 9ofappeared
The analysis in the
the studied current population,
populations showed thatwhich now contains
the Leningrad, all 11 virulence
Astrakhan, and Nizhnigenes. The percent
Novgorod of susceptible
populations strains in the
are rather “potato”
uniform. population
The Astrakhandecreased
“tomato”from 78 to 58%.
population has a unique
The
pattern of analysisgene
virulence of frequencies
the studiedandpopulations
seems to be showed that The
monoclonal. theMoscow
Leningrad, Astrakhan,
population has and Nizhni
Novgorod populations are rather uniform. The Astrakhan “tomato” population has a unique pattern
of virulence gene frequencies and seems to be monoclonal. The Moscow population has the highest
diversity; a high diversity level was also observed for the Mariy El and Kostroma populations that
can be explained by the fact these 3 regions represent potato-growing regions with a high volumes
of imported seed potato.
The data obtained for the studied “potato” and “tomato” populations shows the first ones have a
greater diversity in some parameters, such as the MtDNA haplotype and Pep1/Pep2 loci.
Most of the analyzed populations contained the virulence gene 9, which was not revealed in the
populations, studied in our previous investigation.

Acknowledgments
This study was financially supported by International Science and Technology Center (project #
3714p).
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253
 References
Fry, W.E., Goodwin, S.B., Dyer, A.T., et al. (1993) Historical and recent migrations of Phytophthora
infestans: chronology, pathways, and implications. // Plant Diseases, 77: 653-661.
Gisi, U. and Cohen, Y. (1996) Resistance to phenylamide fungicides: a case study with Phytophthora
infestans involving mating type and race structure. // Annual Review of Phytopathology, 34:
549-572.
Spielman, L.J., Drenth, A., Davidse, L.C., et al. (1991) A second world-wide migration and
population displacement of Phytophthora infestans. // Plant pathology, 40: 422-430.
Sujkovski, L.S., Goodwin, S.B., Dryer, A.T., and Fry, W.E. (1994) Increased genotypic diversity via
migration and possible occurrence of sexual reproduction of Phytophthora infestans in Poland.
// Phytopathology, 84: 201-207.
Day, J.P. and Shattock, R.C. (1997) Aggressiveness and other factors relating to the displacement
of populations of Phytophthora infestans in England and Wales. // European J. of Plant
Pathology, 103: 379-391.
Lebreton, L. and Andrivon, D. (1998) French isolates of Phytophthora infestans from potato and
tomato differ in phenotype and genotype. // European J. of Plant Pathology, 104: 583-594.
Elansky, S.N. (2000) Changes in the Moscow region P. infestans population in 1991-1999. In:
Proceedings of International Conference “Mycology and Cryptogam Botany in Russia”, St.
Petersburg, Komarov Botanical Institute, p. 118 [in Russian].
Goodwin, S.B., Smart, S.D., Sandrock, R.W., et al. (1998) Genetic change within populations
of Phytophthora infestans in the United States and Canada during 1994 to 1996: Role of
migration and recombination. // Phytopathology, 88: 939-949.
Elansky, S.N., Smirnov, A.N., Dyakov, Yu.T., et al. (2001) Genotypic analysis of Russian isolates of
Phytophthora infestans from the Moscow region, Siberia, and Far East. // J. Phytopathology,
149: 605-611.
Hebert, P.D.N. and Beaton, M.J. (1993) Methodologies for allozyme analysis using cellulose acetate
electrophoresis. A practical handbook. – Guelph, Ontario.
Elansky, S.N. and Smirnov, A.N. (2003) Second locus of peptidase as a marker for genetic
investigations of Phytophthora infestans. // Botanica Lithuanica, 9(3): 275-283.
Griffith, G.W. and Shaw, D.S. (1998) Polymorphisms in Phytophthora infestans: four mitochondrial
haplotypes are detected after amplification of DNA from pure cultures or from host lesions.
// Envir. Microbiol., 64, 4007-4014.
Cohen, Y. and Reuveni (1983) Occurrence of metalaxyl-resistant isolates of Phytophthora infestans in
potato fields in Israel. // Phytopathology, 73, 925-927.
Davidse, L.C., Gerritsma, O.C.M., and Velthuis, G.C.M. (1984) A differential basis of antifungal
activity of acylalanine fungicides and structurally related chloroacetanilide herbicides in
Phytophthora megasperma f.sp. medicaginis. // Pesticide Biochem.Physiol., 21, 301-308.
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