Purple coneflower viruses: species diversity and harmfulness
Viral diseases became an actual problem in medicinal plants cultivation. The number of viruses known to infect purple coneflower increased significantly in the last years in many countries. However, there is no any review about the viral diseases of this valuable medicinal crop. Therefore, the aim o...
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irk-123456789-1524292019-06-12T01:26:49Z Purple coneflower viruses: species diversity and harmfulness Dunich, A.A. Mishchenko, L.T. Reviews Viral diseases became an actual problem in medicinal plants cultivation. The number of viruses known to infect purple coneflower increased significantly in the last years in many countries. However, there is no any review about the viral diseases of this valuable medicinal crop. Therefore, the aim of this article is to summarize the main information about the viruses affecting purple coneflower plants (Echinacea purpurea L. Moench.). An analysis of the literature data showed that purple coneflower could be infected by 10 viruses. These viruses belong to the families Bromoviridae, Bunyaviridae, Secoviridae, Potyviridae, Virgaviridae, and almost all of them are considered to be highly harmful plant viruses. Additionally, four of them (TMV, TSWV, CMV, PVY) are in the top 10 of the most economically important plant viruses in the world and occupy the first places. Such distribution and harmfulness of these viruses are explained by a wide range of sensitive host-plants, wild plants and weeds – reservoirs of an infection, and also a large number of vectors. The data from a few countries show that the viral diseases of purple coneflower are becoming more severe from year to year. The appearance of new viruses is registered on coneflower every year that complicates prognosis and risk estimation of epiphytoties in these regions which, for example, were revealed in Bulgaria, Lithuania and Ukraine. This review presents the detailed symptoms of the viral diseases in purple coneflower, the main properties of each virus and data about their harmful effect on the quality of raw material (the concentration of biologically active substances and heavy metals in plants). У світі все більш гострою проблемою при промисловому вирощуванні лікарських рослин стають вірусні захворювання. Встановлено, що в останні роки в багатьох країнах число вірусів, що уражують ехінацею значно зросла. Незважаючи на це, науковцям бракує цілісного огляду світової наукової літератури про вірусні захворювання, які інфікують дану цінну лікарську культуру. Метою роботи є зведення основної інформації про віруси, що заражають ехінацею пурпурову в світі. Аналіз літератури показав, що ехінацея пурпурова уражається 10 вірусами з сімейств Bromoviridae, Bunyaviridae, Secoviridae, Potyviridae, Virgaviridae, практично всі вони визначені шкодочинними фітовірусами. Крім того, чотири серед них (TMV, TSWV, CMV, PVY) займають перші позиції в 10-ці найбільш науково і економічно значущих вірусів рослин в світі. Таке поширення та шкодочинність цих вірусів пояснюється широким колом чутливих рослин-господарів, диких рослин і бур'янів – резервантом інфекції, а також великою кількістю векторів. Дослідження вірусологів з декількох країн показують, що симптоматика вірусних інфекцій ехінацеї з роками стає більш складною. Щорічно діагностуються нові віруси ехінацеї, що ускладнює прогнозування та оцінку ризику появи епіфітотій в певних регіонах, як наприклад, це було зареєстровано в Литві, Україні та Болгарії. У статті детально представлені симптоми вірусних хвороб ехінацеї, основні властивості кожного вірусу і дані про їхній вплив на концентрацію біологічно активних речовин і важких металів рослин. В мире все более острой проблемой при промышленном выращивании лекарственных растений становятся вирусные заболевания. Установлено, что во многих странах число вирусов, поражающих эхинацею в последние годы значительно возросло. Несмотря на это, цельный обзор мировой научной литературы о вирусных заболеваниях, инфицирующих эту ценную лекарственную культуру, отсутствует. Целью работы является суммировать основную информацию о вирусах, заражающих эхинацею пурпурную в мире. Анализ литературы показал, что эхинацея пурпурная поражается 10 вирусами из семейств Bromoviridae, Bunyaviridae, Secoviridae, Potyviridae, Virgaviridae. Практически все они определены вредоносными фитовирусами. Кроме этого, четыре среди них (TMV, TSWV, CMV, PVY) занимают первые позиции в 10-ке наиболее научно и экономически значимых вирусов растений в мире. Такое распространение и вредоносность этих вирусов объясняется широким кругом чувствительных растений-хозяев, диких растений и сорняков – резервантов инфекции, а также большим количеством векторов. Исследования вирусологов из нескольких стран показывают, что симптоматика вирусных инфекций эхинацеи с годами становится более сложной. Ежегодно диагностируются новые вирусы эхинацеи, что усложняет прогнозирование и оценку риска появления эпифитотий в определенных регионах, как например, это было зарегистрировано в Литве, Украине и Болгарии. В статье детально представлены симптомы вирусных болезней эхинацеи, основные свойства каждого вируса и данные о их влиянии на концентрацию биологически активных веществ и тяжелых металлов растениях. 2015 Article Purple coneflower viruses: species diversity and harmfulness / A.A. Dunich, L.T. Mishchenko // Biopolymers and Cell. — 2015. — Т. 31, № 1. — С. 15-28. — Бібліогр.: 77 назв. — англ. 0233-7657 DOI: http://dx.doi.org/10.7124/bc.0008C8 http://dspace.nbuv.gov.ua/handle/123456789/152429 578.3+633.88 en |
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Reviews Reviews Dunich, A.A. Mishchenko, L.T. Purple coneflower viruses: species diversity and harmfulness |
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Viral diseases became an actual problem in medicinal plants cultivation. The number of viruses known to infect purple coneflower increased significantly in the last years in many countries. However, there is no any review about the viral diseases of this valuable medicinal crop. Therefore, the aim of this article is to summarize the main information about the viruses affecting purple coneflower plants (Echinacea purpurea L. Moench.). An analysis of the literature data showed that purple coneflower could be infected by 10 viruses. These viruses belong to the families Bromoviridae, Bunyaviridae, Secoviridae, Potyviridae, Virgaviridae, and almost all of them are considered to be highly harmful plant viruses. Additionally, four of them (TMV, TSWV, CMV, PVY) are in the top 10 of the most economically important plant viruses in the world and occupy the first places. Such distribution and harmfulness of these viruses are explained by a wide range of sensitive host-plants, wild plants and weeds – reservoirs of an infection, and also a large number of vectors. The data from a few countries show that the viral diseases of purple coneflower are becoming more severe from year to year. The appearance of new viruses is registered on coneflower every year that complicates prognosis and risk estimation of epiphytoties in these regions which, for example, were revealed in Bulgaria, Lithuania and Ukraine. This review presents the detailed symptoms of the viral diseases in purple coneflower, the main properties of each virus and data about their harmful effect on the quality of raw material (the concentration of biologically active substances and heavy metals in plants). |
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Article |
| author |
Dunich, A.A. Mishchenko, L.T. |
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Dunich, A.A. Mishchenko, L.T. |
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Dunich, A.A. |
| title |
Purple coneflower viruses: species diversity and harmfulness |
| title_short |
Purple coneflower viruses: species diversity and harmfulness |
| title_full |
Purple coneflower viruses: species diversity and harmfulness |
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Purple coneflower viruses: species diversity and harmfulness |
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Purple coneflower viruses: species diversity and harmfulness |
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purple coneflower viruses: species diversity and harmfulness |
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2015 |
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http://dspace.nbuv.gov.ua/handle/123456789/152429 |
| citation_txt |
Purple coneflower viruses: species diversity and harmfulness / A.A. Dunich, L.T. Mishchenko // Biopolymers and Cell. — 2015. — Т. 31, № 1. — С. 15-28. — Бібліогр.: 77 назв. — англ. |
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15
UDC 578.3+633.88
Purple conefl ower viruses: species diversity and harmfulness
A. A. Dunich, L. T. Mishchenko
Educational and Scientifi c Centre «Institute of Biology»,
Taras Shevchenko National University of Kyiv
64/13, Volodymyrska Str., Kyiv, Ukraine, 01601
korenevochka@mail.ru
Viral diseases became an actual problem in medicinal plants cultivation. The number of viruses known to
infect purple conefl ower increased signifi cantly in the last years in many countries. However, there is no any
review about the viral diseases of this valuable medicinal crop. Therefore, the aim of this article is to sum-
marize the main information about the viruses affecting purple conefl ower plants (Echinacea purpurea L.
Moench.). An analysis of the literature data showed that purple conefl ower could be infected by 10 viruses.
These viruses belong to the families Bromoviridae, Bunyaviridae, Secoviridae, Potyviridae, Vir ga vi ri dae,
and almost all of them are considered to be highly harmful plant viruses. Additionally, four of them (TMV,
TSWV, CMV, PVY) are in the top 10 of the most economically important plant viruses in the world and
occupy the fi rst places. The data from a few countries show that the viral diseases of purple conefl ower are
becoming more severe from year to year. The appearance of new viruses is registered on conefl ower every
year that complicates prognosis and risk estimation of epiphytoties in these regions which, for example,
were revealed in Bulgaria, Lithuania and Ukraine. This review presents the detailed symptoms of the viral
diseases in purple conefl ower, the main properties of each virus and data about their harmful effect on the
plant metabolism and on the quality of raw material (the concentration of biologically active substances and
heavy metals in plants).
K e y w o r d s: purple conefl ower, plant viruses, species diversity, biologically active substances, hea vy
metals.
Introduction
Nowadays demand for herbal medicines is constant-
ly increasing. Purple conefl ower (Echinacea purpu-
rea (L.) Moench.) is an important medicinal plant.
This plant is used for treating more than 70 disorders
in humans and is a component of approximately 300
herbal medicines [1]. Besides that, it is also used as
ornamental, melliferous and essential oil plant. Pur-
ple conefl ower is an herbaceous perennial plant of
Asteraceae family. It is native to the eastern and so-
uthern USA [2].
This herb is cultivated in the north-western sta tes
of America, in western Canada, Australia, New
Zealand, southern America, Europe and also was in-
troduced in Ukraine. The diffi culty of its cul tivating
is connected with the condition requirements for the
plant. Unlike the natural phytocenosis, during ac-
commodating to agrocenosis conditions the herbs
are becoming more wasted and susceptible to plant
pathogens of different etiology. Besides that, purple
conefl ower is a perennial plant that causes accumu-
lation of pathogens, including viruses.
Regardless of the value of the purple conefl ower in
different areas, there is no review on its viruses all over
the world. So, in this article we present a review that
summarizes the main information about the viruses af-
fecting purple conefl ower plants all around the world.
There are data about diseases of purple conefl ow-
er caused by 10 viruses (Tabl. 1).
© 2015 A. A. Dunich et al.; Published by the Institute of Molecular Biology and Genetics, NAS of Ukraine on behalf of Biopolymers and Cell.
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/),
which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited
ISSN 0233-7657
Biopolymers and Cell. 2015. Vol. 31. N 1. P. 15–28
doi: http://dx.doi.org/10.7124/bc.0008C8
16
A. A. Dunich, L. T. Mishchenko
Viruses of Bromoviridae family
Alfalfa mosaic virus, AMV. Alfalfa mosaic virus is a
member of Alfamovirus genus, Bromoviridae family.
AMV has a wide host range. This virus can naturally
infect many herbaceous and some woody plant hosts
(150 species of 22 families) and is transmissible to over
430 species of 51 dicotyledonous families [25]. AMV
can cause various mosaics, mottles, and malformations
in alfalfa (Medicago sativa), yellowing of lea ves in pea
(Pisum sativum), calico and tuber necrosis in potato
(Solanum tuberosum), and various symptoms in tobac-
co (Nicotiana tabacum) [26, 27]. Sixteen species of
aphids, including Myzus persicae, can tran smit AMV
in a non-persistent manner [28]. AMV can be also
transmitted by potato pollen and by seed [29].
In 2008–2009, Alfalfa mosaic virus was detected
in Bulgaria in 55.5 % of tested purple conefl ower
samples [3, 4]. AMV caused spotting and dwarfi ng
of the purple conefl ower leaves. The virus was de-
tected in co-infection with TMV and CMV. The im-
pact of the disease caused by CMV, AMV and TMV
on the conefl ower yield was also determined. Infected
herbs had lower weight and number of leaves, ra-
cemes and produced seeds half as much.
Cucumber mosaic virus, CMV
Cucumber mosaic virus causes substantial agronomic
yield losses in many crops in the world, probably has
one of the broadest host range among plant viruses.
CMV is a type member of the genus Cu cu mo virus,
family Bromoviridae and infects more than 1,000 plant
Table 1. Viruses that infect purple conefl ower in the world
Virus name Acronym Genus, family
Detection
on conefl ower,
country
References
Alfalfa mosaic virus AMV Alfamovirus, Bromoviridae Bulgaria [3, 4]
Broad bean wilt virus 2 BBWV-2 Fabavirus, Secoviridae China [5]
Cucumber mosaic virus CMV Cucumovirus, Bromoviridae Germany [6]
New Zealand [7, 8]
Japan [9]
Italy [10–12]
USA [13]
Hungary [15]
Belarus [16]
China [22]
Bulgaria [3,4]
Ukraine [17]
Impatiens necrotic spot virus INSV Tospovirus, Bunyaviridae Lithuania [18]
Potato virus Y PVY Potyvirus, Potyviridae Hungary [14]
Bulgaria [3,4]
Ukraine [19]
Tobacco mosaic virus TMV Tobamovirus, Virgaviridae Bulgaria [4]
Hungary [14]
Tobacco rattle virus TRV Tobravirus, Virgaviridae Lithuania [18, 20]
Tobacco ringspot virus TRSV Nepovirus, Secoviridae Lithuania [20, 21]
Tomato ringspot virus ToRSV Nepovirus, Secoviridae Lithuania [22]
Tomato spotted wilt virus TSWV Tospovirus, Bunyaviridae Lithuania [18]
Hungary [14]
Bulgaria [4, 23]
Ukraine [24]
17
Purple conefl ower viruses: species diversity and harmfulness
species from more than 86 families, including mono-
cots and dicots [25]. The host range includes different
crops and numerous wild plants, which are important
for the annual persistence of the virus in the open fi eld.
CMV is transmitted by aphides (Aphididae; mo re
than 60 species, including Acyrthosiphon pisum, Ap his
craccivora and Myzus persicae) in non-persistent man-
ner, by mechanical inoculation and seeds [25, 30].
The earliest report on CMV affecting purple cone-
fl ower was in 1964 in Germany [6]. CMV-infected
purple conefl ower was revealed in Belarus and re-
ported as a weedy host for the virus [15], New
Zealand with the symptoms of yellow mottling on
the leaves [7, 8], and also in the US, where the dis-
ease was accompanied with ringspots and leaf defor-
mation [13].
In 1993, CMV disease of Echinacea purpurea was
observed in Japan [9]. Conefl ower plants showed
mosaic symptoms in the leaves.
Yamamoto et al. studied a host range of an isolate
of CMV (39 plant species were inoculated) [9]. It
was demonstrated that 18 species including Echi-
nacea purpurea, Lycopersicon esculentum, Petunia
hybrid, Capsicum annum, Pisum sativum, Vicia fa-
ba, Cucumis sativus, Nicotiana tabacum, N. gluti-
nosa were systemically infected by CMV, but the
virus could not be readily transmitted to Cucumis sa-
tivus. Eighteen species such as: Citrullus lanatus,
Cucumis melo, C. melo (makuwa), C. melo (cono-
mon), Cucurbita maxima, C. moschata, C. pepo, Vig na
ses quipedalis, Chenopodium quinoa, Che nopodium
amaranticolor were locally infected. It was the fi rst
report about the viral disease of purple conefl ower in
Japan and the authors proposed to name this disease
as ‘Echinacea mosaic disease’ [9].
In Italy, CMV has been detected in the plants of
purple conefl ower for years. The disease has the sym-
ptoms of yellow mosaic, variegation on the leaves
which were often malformed and with bullas. The
petals showed contractions, deformations and mot-
tling [10, 11]. In 2009, CMV-infected plants showed
stunting; leaves with yellow mosaics, ring, line pat-
terns and malformations; small fl owers with pale
stripes on red petals [12]. Mo reover, Italian scien-
tists have studied genetic modifi cations of the CMV
isolates, taken from aromatic, offi cinal and orna-
mental plants in the Northern Italy. They made a se-
quence analysis of the movement protein gene of
CMV isolate from purple conefl ower and its com-
parison with the isolates from other countries [31]. A
phylogenetic analysis showed that CMV isolate from
Echinacea purpurea from Italy (EU 432180) be-
longed to the subgroup IA and was genetically closer
to the Italian isolate CMV-LAV (EU432182) from
Lavandula offi cinalis and Hungarian NS isolate from
Nicotiana glutinosa (AJ511990). The comparison of
nonsynonymous and synonymous substitutions sug-
gested that 30 % of amino acid sites were under neg-
ative selection and only one was under positive se-
lection. Phylogenetic, nucleotide diversity and ge-
netic differentiation analyses suggested that long-
distance migration plays a role in the evolution and
determination of the genetic structure and diversity
of CMV in northern Italy and other regions.
CMV was also revealed in Hungary in mixed in-
fection with PVY, TMV і TSWV [14].
The disease of purple conefl ower with the symp-
toms of yellow mosaics on the leaves was also re-
vealed in China. On the basis of virion structure, the
biological and serological properties were deter-
mined that were caused by CMV [16]. The nucle-
otide and amino acid sequence analysis of the RNA
2-1-1 fragments showed the 94.5 and 98.1 % identity
with the standard isolates of Fny subgroup as well as
77.7 and 82.5 % identity with the standard isolate Q
from the subgroup II [32].
In Bulgaria, 45.5 % of purple conefl ower plants
with viral symptoms were determined as infected by
CMV [4]. The symptoms were light-green spots on
the leaves with curly laminas due to the different
growth of pale green and dark green areas of the
leaves (Fig. 1).
Dikova et al. showed that economically important
viruses for the E. purpurea cultivation in Bulgaria
are those, transmitted by aphids: AMV and CMV as
well as the mechanically transmitted TMV [4]. These
three viruses were widespread in over 45 % of plants
and caused the symptoms of spotting and even of
mosaic on the leaves. It is interesting to mention that
mixed infection by CMV, AMV and TMV caused
18
A. A. Dunich, L. T. Mishchenko
Fig. 2. E. purpurea infected by CMV, AMV and TMV in Bulgaria: A — spotting on the leaves; B — dwarfed leaves and stunting [4]
Fig. 3. Echinacea purpurea infected by CMV in Ukraine: on each photo at the left – healthy plant, at the right – virus infected plant [17, 33]
Fig. 4. Symptoms of TSWV infection on purple conefl ower in Ukraine: А, B – phase of budding-initial blossoming; C – end of blos-
soming [19, 24, 61]
A
A
A
B
B
B
C
C
Fig. 1. Symptoms of mosaics on Echinacea purpurea leaves cau sed by Cucumber mosaic virus in Bulgaria (at the left – a symptomless leaf) [4]
other symptoms – spotting and dwarfed conefl ower
leaves (Fig. 2).
Dikova and co-authors revealed that sprouts,
containing viruses, grew from the purple cone-
fl ower roots, infected during the fi rst year of two-
and three- year old conefl ower plantations showed
the highest percentage of infection with spotted
plants and dwarfi ng leaves and stems [3, 4]. They
showed that the vector of CMV could be aphids
Aphis gossypii Glover (cotton aphid) which were
found in the trial fi elds with E. purpurea during
April, May and June. These aphids were not ob-
served at later stages of vegetation of E. purpurea
plants in July and August.
In 2006–2009, epiphytoty of CMV on fi eld-grown
purple conefl ower was registered in Ukraine [17, 33].
Monitoring of the plantations showed that 75 % of the
plants were with the symptoms of viral infection. The
most typical symptom was chlorosis on the leaves ac-
companied by their size reduction (Fig. 3, A).
19
Purple conefl ower viruses: species diversity and harmfulness
It is noteworthy that these symptoms were revealed
among the herbs of the 1st year of cultivation as well as
among those of the 2nd–5thyears during both regenera-
tion and blossoming periods. Also rolling, rasp-shape
and leaf deformation were revealed on the 2nd year
plants (Fig. 3, B). In limited number of the cases, the
plant stunting and the stunting of the root system have
been observed (Fig. 3, C). Generally, it was observed
that the virus symptoms were getting more pronounced
during the last years.
The spherical particles were revealed in the purple
conefl ower plants 29.5 ± 0.5 nm in diameter [17, 34].
The virus identifi cation carried out by ELISA and
the study of biological, physical and chemical prop-
erties of the virus proved that the plants were infec-
ted by CMV [17]. It was the fi rst report about in-
fection of purple conefl ower by CMV in Ukraine.
Taking into consideration the data about species
diversity of the viruses infecting purple conefl ower
in the world and those widely spread in Ukraine, the
samples were also tested as to the presence of other
viruses. The results of ELISA test indicated that in
purple conefl ower there were no antigens of the fol-
lowing viruses: TMV, AMV, TSWV, INSV, TRV,
PVY, PVS, PVX, PVM, TAV, ArMV, TuMV, PePMV,
WMV-2, and CGMMV [17].
Viruses of Bunyaviridae family
Tomato spotted wilt virus, TSWV, and Impatiens ne-
c rotic spot virus, INSV. TSWV and INSV are the
members of genus Tospovirus. Initially INSV was con-
sidered to be another isolate of TSWV, but genetic stud-
ies proved that they are different virus species [35].
Tospoviruses are transmitted in a persistent and propa-
gative manner exclusively by thrips (Thysanopte ra:
Thripidae) [36]. Trans mis sion can also be achie ved
through infected plant sap. INSV and TSWV are
mainly defi ned on the basis of their vector spe cifi city,
host range and symptoms of disease.
TSWV was reported to have a very broad host
range including more than 1100 different species
from more than 80 families [37]. The TSWV infec-
tion is characterized by different symptoms de-
pending on the plant species [38, 39]. TSWV causes
severe diseases in numerous horticultural and agri-
cultural crops. TSWV is the second among the top
10 of scientifi cally and economically important
plant viruses in the world [40]. The distinguishing
feature of this virus is an ability to infect many spe-
cies: from 150announced in 1968 [41] to 650 an-
nounced in 1994 [42] and about 1100 plants from
more than 80 families – in 2003 [37, 43]. The num-
ber of susceptible plants exposed to the risk to be
infected by TSWV is growing nowadays. It can be
explained by some features of epidemiology of this
virus. There are innumerable infected weeds that
serve as reservoirs for primary infection [38]. These
plant sources perpetuate TSWV as important hosts
for thrips vectors and serve as foci for subsequent
movement and infection of susceptible crop plants
[44]. TSWV is transmitted by thrips Frankliniella
occidentalis, F. bispi nosa, F. cephalica, F. gemina.
F. fusca, F. intonsa, F. schultzei, F. se tosus and
Thrips tabaci [25, 36]. TSWV is also transmitted
by grafting; and is not transmitted by contact be-
tween plants, by seed and by pollen.
INSV has a narrower host range than TSWV.
INSV infects more than 648 species including im-
portant horticultural and agricultural crops [25]. Its
name indicates that the main symptom is necrotic
spotting on the leaves. However, it is rather diffi cult
to diagnose INSV as the symptoms can vary depend-
ing on the host and its age. The symptoms caused by
INSV can be easily confused with those of other vi-
ruses, fungi and bacteria or nutrition disorders. It has
been announced many years ago that fl ower thrips
(Frankliniella occidentalis) are the only effective
transmitter of INSV [45]. Mo re over, INSV can be
transmitted by other species Fran kliniella — F. in-
tonsa [46] and F. fusca [47].
In purple conefl ower TSWV was detected in Euro-
pe (Lithuania, Bulgaria, Hungary and Ukraine). In
Lithuania, TSWV on the purple conefl ower was re-
gistered in co-infection with INSV [18].
In 2009–2010, TSWV was also revealed in Bul-
garia on the purple conefl ower plants [23]. The au-
thor described that TSWV caused yellow spotting on
the purple conefl ower leaves. TSWV was revea led
in one plant with the symptoms on leaves typical for
this virus – chlorotic dark red ring spots turning into
20
A. A. Dunich, L. T. Mishchenko
brown necrotic lesions [4, 23]. Thripses were found
on E. purpurea racemes in single cases. The main
species was Frankliniella occidentalis that appeared
in May to July.
In 2012, purple conefl ower disease caused by
TSWV was revealed in Ukraine [24]. The symptoms
of the disease were yellow spots, mosaics and leaf
deformation during all vegetation phases. Light
green mottling on the leaves during budding-initial
blossoming (Fig. 4, A, B) turned into yellow mot-
tling, that at the end of blossoming covered practi-
cally all lamina (Fig. 4, C).
The virions of the TSWV isolate were studied
with the electron microscopy method and were 100 ±
± 20 nm in diameter.
It was also studied whether the plant samples were
infected by INSV taking into consideration that in
Lithuania purple conefl ower was infected with the
complex of TSWV and INSV [18]. The ELISA re-
sults showed the absence of INSV in the conefl ower
plants [24].
Viruses of Potyviridae family
Potato virus Y, PVY. Potato virus Y is a type mem-
ber of the genus Potyvirus of family Potyviridae. It
is common knowledge that this is one of the most
widely spread and economically important pathogen
with a broad range of susceptible plants from differ-
ent families. Besides the potato it can infect tobacco,
tomato, sweet pepper and many others. Nowadays
PVY is the fi fth among top 10 economically and sci-
entifi cally important plant viruses [40]. The virus is
transmitted by a vector in non-persistent manner (ar-
thropods, insects of order Hemip tera, family Aphi di-
dae), by mechanical inoculation, by grafting. Trans-
mitting by seeds and contact is plausible [25, 48].
In 2008, only 3.13 % of purple conefl ower infec-
ted by the PVY+ CMV+AMV complex was reve-
aled in Bulgaria. The symptoms were spotting and
«dwarfed» leaves [3, 4]. In Hungary PVY was found
in the co-infection with CMV, TMV and TSWV [14].
PVY was also detected in Ukraine in purple co-
nefl ower with the symptoms of leaf curling and de-
formation [19]. The samples of 2nd and 3rd years of
cultivation were tested positive by RT-PCR.
Viruses of Secoviridae family,
Comovirinae subfamily
Broad bean wilt virus 2, BBWV-2. BBWV-2 is a
member of Fabavirus genus [49]. BBWV-2 spreads
in African region, Eurasian region, Middle East,
North American region, Pacifi c region, Australia and
China. The virus is transmitted by a vector in a non-
persistent manner; an insect; Acyrthosiphon pisum,
Aphis craccivora, A. faba, A. nasturtii, Mac ro siphum
euphorbiae, M. solanifolii, M. persicae; Aphididae.
The virus is transmitted by mechanical inoculation
and not transmitted by seeds.
In 2010, BBWV-2 was revealed on purple cone-
fl ower plantations in China. The symptoms were ne-
croses, leaf rolling, yellow mosaics, and mosaics in
leaves [5]. One plant sample with mosaics symptoms
was tested positive in ELISA. Li et al. have studied the
range of experimental hosts. Susceptible plant species
for Chinese BBWV-2 isolate were Chenopodium qui-
noa (local lesions in inoculated lea ves, systemic chlo-
rotic mottle in upper leaves, deformation, and apical
necrosis), C. amaranticolor (chlo rotic local lesions,
systemic mosaic and leaf distortion), Nicotiana ben-
thamiana (systemic mosaics), Gomphrena globosa
(local purple spots in inoculated leaves and systemic
infection in upper leaves), Tetragonia expansa (local
lesions, but no symptoms of systemic infection),
Physalis fl oridana (systemic mosaics). No symptoms
were observed on Capsicum annum, Datura stramo-
nium, N. glutinosa, or N. tabacum cv. White Burley.
Compa rison with the sequences of other BBWV-2
isola tes showed that the isolate from conefl ower
(No. JX070674) had approximately 99 % nt identity
(98 % amino acid identity) with Chinese BBWV-2 iso-
late BC from Bupleurum chinense. This was the fi rst
report of BBWV-2 infecting purple conefl ower in
China and, to our knowledge, in the world.
Tobacco ringspot virus, TRSV and Tomato ring-
spot virus, ToRSV. TRSV and ToRSV belong to the
subgroup A of genus Nepovirus and are serologically
related viruses [25]. The nepoviruses have a wide
host range, including both woody and herbaceous
plants. TRSV causes signifi cant disease in gra pe,
soybean, tobacco, blueberry and members of Cu-
21
Purple conefl ower viruses: species diversity and harmfulness
curbitaceae family (melon, cucumber, squash, and
pumpkin). Ma ny other hosts infected naturally were
found, including apple, pepper, cherry, papaya, vari-
ous weeds and mint [50]. TRSV causes systemic
chlorotic or necrotic ringspots, leaf deformation and
stunting. The ToRSV host range is very similar to
that of To bacco ringspot nepovirus, except that it is
much less important for fruit crops than ToRSV.
The nepoviruses are transmitted by the nematode
Xiphinema americanum and other closely related
Xiphinema spp [51]. TRSV is easily transmitted me-
chanically, by seeds and pollen [52, 53]. TRSV and
ToRSV have been reported in Europe, North Ame ri-
ca, Australia, Africa, India, Japan [3] and New Zea-
land [54], Lithuania [55] and Uk raine. TRSV is in-
cluded in the list of quarantined pest [56].
Firstly ToRSV on the purple conefl ower plants was
revealed in Lithuania in 2000 [22] and TRSV in 2006
[21]. In 2003, the mixed infection of ToRSV and TRV
was detected [18]. In 2008, Echinacea purpurea (L.)
Moench exhibiting the symptoms of plant stunting, leaf
malformation, various shaped chlorotic spots, ringspots
was collected in the Botanical Gar den of Vilnius
University. According to the results obtained by the
methods of test-plants, electron microscopy, DAS-
ELISA, and RT-PCR, the purple conefl ower plants
were affected by co-infection of TRSV and TRV [21].
Viruses of Virgaviridae family
Tobacco rattle virus, TRV. Tobacco rattle virus is a
type member of Tobravirus genus, Virgaviridae fa-
mily. For the fi rst time this virus was revealed in 1931 in
Germany on tobacco. TRV is transmitted by a vector
(nematodes; family Trichodoridae; Paratri chodorus
allius, P. anemones, P. christiei, P. nanus, P. pachy-
dermus, P. teres, Trichodorus minor, T. pri mitivus, T.
viruliferus). The virus is transmitted by mechanical
inoculation, grafting and seeds; not tran smitted by
contact between hosts.
Geographically TRV has been found throughout
Europe, New Zealand, North America and Ja pan.
TRV has one of the widest host ranges among all the
plant viruses. Natural infection has been reported in
more than 100 plant species [25]. About 400 species
in more than 50 families can be infected by sap in-
oculation [57]. TRV has continuously been a signifi -
cant potato pathogen causing corky ringspots in po-
tato tubers, which renders the crop unmarketable
[58]. Additionally, infection by TRV may cause a
loss of vigor and yield in tomato, tobacco, sugar
beet, spinach, artichoke, celery, pepper and lettuce.
In 2003, TRV was detected in the purple cone-
fl ower in co-infection with ToRSV and in 2008 with
TRSV in Lithuania [18, 20]. This is the only report
about infecting purple conefl ower plants by TRV in
the world.
Tobacco mosaic virus, TMV is the type member
of genus Tobamovirus. TMV is thermostable, the
thermal inactivation point is 95 C. It is extremely
persistent in an external environment. The virus is
not transmitted by a vector. Virus is transmitted by
mechanical inoculation, grafting, contact between
hosts, by seeds (occasionally transmitted through the
testa, but not through the embryo), not transmitted
by pollen [25].
TMV has a wide host range (more than 350 species),
which includes various vegetables (beet, pepper, egg-
plant, potato, spinach, tomato), agricultural crops (to-
bacco, beans, grapes, apple-tree), de corative plants (pe-
tunia, phloxes, zinnia), annu al and perennial weeds. It is
one of the most contagious plant viruses. The sources
of infection are plant residues, seeds, and soil in which
the virus does not lose viability for over 22 months. The
sym ptoms vary depending on the virus strain, species,
phase of infection, environmental conditions. A mixed
infection of Tobacco mosaic virus with other viruses
(Potato virus X, Potato virus Y, Cu cum ber mosaic
virus) causes brown wide and narrow stripes on fru-
its, leaves, stems and petioles which die off in course
of time. TMV is distributed in Eurasia, revealed in
Argentina, Australia, Germany, France, Hungary, Ice-
land, India, Italy, Japan, Pe ru, Kenya, Spain, USA,
and Great Britain [25].
TMV in purple conefl ower plants was revealed in
Hungary in 2006 [14] and in Bulgaria [4], where it
was observed in a co-infection with AMV and CMV.
TMV caused the mosaics symptoms and spotting
and is considered to be one of the most economically
important viruses infecting purple conefl ower (Fig. 2).
Dikova et al. found that the virus infected conefl ow-
22
A. A. Dunich, L. T. Mishchenko
er plants were lower, with smaller number of leaves
and racemes in comparison with the control plants
[4]. The differences in the sizes of organs affected
the yield. So, the yield of leaf mass (herba) from the
virus infected plants was twice to several times low-
er in comparison with the symptomless samples. It
was noted that more than half of the purple cone-
fl ower plants died over a period of three years. Such
perishing was due to different factors – viruses, phy-
toplasma pathogens and environmental conditions
(dry soil, high or low temperature). In order to mini-
mize the damage, caused by viruses, the purple cone-
fl ower fi elds should be isolated from the vegetable
and fl ower plants [4].
Unidentifi ed viruses
Rod-shaped virions 40 ± 5, 70 ± 5 and 130 ± 30 nm
long, 17–20 nm wide were revealed in the leaves of
purple conefl ower cultivated in Ukraine (Fig. 5).
Viruses, presented in Fig. 5, are rod-shaped with
distinct axial canal. Morphologically they are similar
to the viruses of Virgaviridae family, which includes
six genera: Hordeivirus, Furovirus, Pec lu vi rus, Po mo-
virus, Tobamovirus, and Tobravirus. How ever, among
mentioned genera, the fi rst four have a very narrow
host range. The described viruses can not be consid-
ered as tobamoviruses as the length of virions differs
substantially. Morpho lo gically they are closer to to-
braviruses. Additionally, the infecting purple cone-
fl ower plants by Tobacco rattle virus which belongs
to this family was registered in Lithuania [18, 20].
Tobraviruses are transmitted by the nematodes Tri-
chodorus and Parat richodorus (Trichodoridae), which
are widespread in Ukraine [59]. However, the results
of ELISA tests indicated that this rod-shaped virus is
not the TRV (unpublished data). Maybe, purple
conefl ower in Ukraine was affected by other mem-
bers of the genus Tobravirus: Pea early browning
virus, PEBV or Pepper ringspot virus, PepRSV. How-
ever, testing conefl ower plants on the antigens of
PEBV and PepRSV was not conducted. There are
the data about infecting atypical plants by hordeivi-
ruses which is known to have a very narrow host
range which includes mainly cereals. Beczner et al.
noted that in Hungary the member of Hordeivirus
genus Lychnis ringspot virus (LRSV-M) infected
mint plants (Mentha longifolia Huds) [60]. The virus
is transmitted mechanically, by seeds and pollen.
Taking this into consideration, we should not elimi-
nate the possibility that purple conefl ower can be
infected by hordeiviruses in Ukraine.
Effect of plant viruses
on the general links of plants
metabolism
For today, despite numerous researches devoted to
the study of viral diseases of plants, the mechanisms
of development of the pathological process remain
unexposed and, at the same time, actual for many sci-
entists. Study on the effects, caused by a viral infec-
tion, at the level of cells and their compartments as
well as at the organism level is important for under-
standing the mechanisms of co-operation of the virus
with a plant. The research of infl uence of viral infec-
tion on the basic links of metabolism is of special in-
terest as they provide the plant with plastic and power
substances that are necessary for normal development
and vital functions.
Many scientists investigate an infl uence of viruses
on the photosynthetic apparatus of plants, in fact the
level of photosynthesis is one of basic indexes of the
plant normal development and productivity, and
thus, the plant: resistance to the infections [17, 61–
63]. They revealed a substantial reduction in the
concentration of both chlorophylls a and b as well as
of carotenoids in the leaves of Lophanthus anisatus,
Arctium lappa and Echina cea purpurea under viral
infection. The chlorophyll a content in the leaves
of virus infected Lophanthus anisatus plants was by
35 % lower, in comparison with the healthy stan-
dards. This tendency was observed also at research
of concentration of chlorophyll b, that in sick plants
diminished three times (by 31.4 % in comparison
with the control). The carotenoids concentration di-
minished by 32 % in the virus infected giant hyssop
plants [64]. A substantial decline in the photosyn-
thetic pigments content under the action of viral in-
fection was found for the burdock plants too. Con-
centration of chlorophyll a was less by 69.7 % com-
pared with the control, chlorophyll b – by 73.0 %,
23
Purple conefl ower viruses: species diversity and harmfulness
sum of carotenoids – by 44.6 %. In leaves of the
CMV-infected Echinacea purpurea plants content of
chlorophyll a was by 50.5 % lower, than in healthy
plants, and content of chlorophyll b and sum of ca-
rotenoids – by 68.9 % and 41.3 %, accordingly [61].
It is necessary to mark that the most substantial de-
cline in the content of these pigments was revealed
in purple conefl ower plants that testify about high
harmfulness of the virus.
Viruses can infl uence the metabolism of plants
differently, including the content of carbohydrates
in leaves. Some viruses have insuffi cient effect on
carbohydrates in leaves, whereas others can change
both speed of synthesis and speed of their move-
ment in a plant. It was revealed that in leaves of the
infected ginseng plants the content of saccharides
was higher compared with the healthy plants:
monosaccharides – by 26 %, sucrose – by 102 %,
sum of sugars – by 57 % [17]. One of suggestions
of a reason of glucose, fructose and sucrose accu-
mulation in the leaves of plants infected by a virus
was the violation of their outfl ow from phloem that
conduces to the development of gummosis. Later
Watson M.A. and Watson D.J. made a conclusion
that the carbohydrates accumulation in leaves of
the sugar beet staggered by the Beet yellow virus is
conditioned not by blocking the carbohydrates out-
fl ow from leaves but by the infl uence of the virus
on the cells of leaf cells and, maybe, related to the
changes in the activity of the enzyme systems that
control the interconversion of different forms of
carbohydrates [65]. It was shown that viral infec-
tion conduces to the increase in quantitative content
of carbohydrates in leaves and stems of ginseng plants
(Panax ginseng C.A. Meyer): mo nosaccharides – on
26 %, sucrose – on 102 %, sum of saccharides – on
57 % in comparison with the uninfected plants [17].
Unlike ginseng, in purple conefl ower plants infect-
ed with CMV there was registered decline of con-
tent of carbohydrates – monosaccharides – by 58.8 %,
sum of saccharides – by 7.7 % [17]. Such contro-
versial data on the carbohydrates concentration for
these two herbs (conefl ower, ginseng) at a viral in-
fection can be explained by the fact that ginseng
was infected by a less virulent virus, than culture of
Echinacea purpurea. Probably, in the last case a
tendency to the accumulation of sugars is counter-
balanced by more active infl uence of viruses on the
photosynthetic apparatus of a plant.
Danger and harmfulness of viral diseases increas-
es due to some factors. Firstly, a viral infection nega-
tively affects a height and development of plants,
represses the process of forming productive stems
and reproductive organs. Secondly, a viral disease
makes weakened plants more vulnerable to other
pathogens, in particular, by fungi that cause the root
rot. Thirdly, a viral infection often passes and spread
by vectors, especially by insects, that is a diffi cult
and ecologically dangerous challenge.
Effect of plant viruses
on the raw material quality
Except a signifi cant effect of viruses on the yield of
purple conefl ower and other medicinal plants [4, 17],
it was proved that these pathogens reduce the content
of biologically active substances in medicinal plants.
In Italy, virologists showed that CMV reduced the
content of some lipophilic compounds in the purple
conefl ower plants [10, 11, 66]. The infected materials
from Echinacea purpurea afforded a lower oil con-
tent and signifi cant quantitative variations in the oil
composition when the plants were infected by CMV
[67]. It was shown that the CMV infection led to a
Fig. 5. Rod-shaped virions from Echinacea purpurea cultivated
in Ukraine [19, 61]
24
A. A. Dunich, L. T. Mishchenko
decrease in the concentration of hydrocinnamic acids
sum, caftaric and cichoric acids sum in conefl ower
plants, sometimes to the level below the norms regu-
lated by the State Pharmacopoeia [61, 68, 69]. The
polysaccharides content signifi cantly reduced, too
[17, 69]. Pellati with colleagues have studied the con-
tent of the secondary metabolites, such as coffee acid
derivatives, alcamides and essential oil in purple
conefl ower affected by CMV and phytoplasma
16SrIX-C [12]. It was noted, that in the plants infect-
ed by both pathogens, the concentration of cichoric
acid substantially diminished. Viral infection reduces
the content of alkamides and some components of es-
sential oil [12]. Regarding the main alkamide, dode-
ca-2E,4E,8Z,10E / Z-tet rae no ic acid isobutylamide, a
signifi cant decrease in the content of this secondary
metabolite was observed in virus-infected plants in
comparison with healthy plants, whereas in the phy-
toplasma-infected sample the variation of this sec-
ondary metabolite was not appreciable.
Indisputable negative CMV infl uence on the qual-
ity of raw material was also revealed for some other
medicinal plants. For example, it was shown that the
CMV infection reduced signifi cantly the quantitative
yield of essential oil in Agastache anethiodora and
caused changes in the relative composition of the
main components: pulegone, menthone, iso-men-
thone, methyl chavicole and limonene [70].
Negative infl uence on the raw material quality is
also shown for some other viruses. It was discovered
that AMV reduces the essential oil secretion in the
lavender plants [71], Peppermint stunt virus – in the
mint plants [72] and BBWV-1 reduces quantitative
and qualitative content of essential oil in the Thymus
vulgaris [73] and Salvia sclarea plants [74, 75]. A
decrease of the basic BAS in the infected plants is dis-
covered also for ginseng (Panax ginseng C.A. Meyer).
It was shown that the content of triterpene glycosides
(saponins) in the roots of infected ginseng was 5.6 %,
i.e. by 2.0 % less than in the healthy plants [17]. Al-
though such raw material remains suitable for phar-
macology, however, such substantial worsening of the
raw material quality can have catastrophic conse-
quences taking into account diffi culty of cultivating
this crop in Ukraine.
Except the infl uence of viruses on therapeutically
active substances in the plants it was shown that these
pathogens worsened the quality of raw material for
drug production. So, upon the action of viral infection
it was revealed the accumulation of some heavy met-
als in medicinal plants in amo unts which exceed max-
imum allowable concentrations (MAC) in food prod-
ucts. It is necessary to notice that the content of high-
toxic elements (As, V, Sb, Cr, Fe) in the TSWV-
infected purple conefl ower plants exceeded MAC in
1.2, 7, 2.3, 2.5 and 3.4 times respectively, unlike the
healthy plants, the concentration of these metals [in
which] was within the limits of norm [24, 76].
In the virus infected Lophanthus anisatus plants the
concentration of Pb, As, Cu, Zn, V, Ni, Fe and Al ex-
ceeded MAC in 1.1, 4.9, 1.7, 3.5, 8.4, 5.8, 7.3, 13.2
times respectively [77]. There is no clear information
about the passing of heavy metals to the medical
forms. That is why viruses can cause a serious prob-
lems in the production of high-quality me dical raw ma-
terial from infected medicinal plants.
Discussion
The review of world scientifi c literature concerning
viruses, which infect purple conefl ower crops in dif-
ferent countries, showed that almost all of them are
considered to be highly harmful and economically
important plant viruses. Additionally, four of them
(TMV, TSWV, CMV, PVY) occupy the fi rst places
in top 10 of the most economically important plant
viruses in the world [40]. Such prevalence and harm-
fulness of these viruses are explained by a wide range
of susceptible host plants, wild plants-reservoirs of
the infection, and a variety of vector species.
Thus, the analysis of literature showed that all vi-
ruses, registered on purple conefl ower, are po ly pha-
gous. Additionally, the number of viruses known to in-
fect purple conefl ower has increased signifi cantly in the
last years. Every year new viruses appear in the same
agrocoenosis, that was revealed, for example, in Bul-
garia, Lithuania and Ukraine [4, 17, 18, 21, 22, 24, 34,
62, 69]. In fact, it complicates the prognosis and risk
estimation of epiphytoties emergence in these regions.
Detection and circulation of such harmful viruses
on commercial plantations of purple conefl ower re-
25
Purple conefl ower viruses: species diversity and harmfulness
quire permanent virological inspections. Cultivation
of medicinal plants needs the safety management. In
case with medicinal plants using of chemical sub-
stances against viruses and their vectors is unaccept-
able. That is why well-timed detection of plant vi-
ruses and information about symptoms of disease,
main properties of these pathogens is almost the only
method of the prevention of viral infections. Strict
control, well-timed diagnostics and disease manage-
ment of medicinal plants must be applied by the es-
tablishments cultivating them. It would reduce the
risk of viruses migration into new environments with
new vectors that could lead to serious diseases not
only of purple conefl ower, but also other horticul-
tural or ornamental crops.
A signifi cant reduces of the BAS and accumula-
tion of toxic metals in different medicinal plants un-
der viral infection have been shown in many coun-
tries. Today one of the general principles of the mod-
ern phytotherapy is medicines safety. That is why
much attention of WHO and national Pharmacopoeias
is paid to the control of pollution of medicinal raw
materials with pesticides, radionuclides, heavy met-
als, mycotoxins. So, we think that the information
about such harmfulness of plant viruses must attract
an attention of the drugs producers, creators of the
pha rmacopoeia papers, state standards and specifi -
cations which regulate the quality of medicinal raw
material. Virus detection should be included to the
worksheets of medicinal crops and to the State
Pharmacopoeias as one of key principles for obtain-
ing high yield and high-quality raw material.
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А. А. Дуніч, Л. Т. Міщенко
Віруси ехінацеї пурпурової:
видове різноманіття та шкодо чинність
У світі все більш гострою проблемою при промисловому
вирощуванні лікарських рослин стають вірусні захворю-
вання. Встановлено, що в останні роки в багатьох країнах
число вірусів, що уражують ехінацею значно зросла.
Незважаючи на це, науковцям бракує цілісного огляду сві-
тової наукової літератури про вірусні захворювання, які ін-
фікують дану цінну лікарську культуру. Метою роботи є
зведення основної інформації про віруси, що заражають
ехінацею пурпурову в світі. Аналіз літератури показав, що
ехінацея пурпурова уражається 10 вірусами з сімейств
Bromoviridae, Bunyaviridae, Secoviridae, Potyviridae,
28
A. A. Dunich, L. T. Mishchenko
Virgaviridae, практично всі вони визначені шкодочинними
фітовірусами. Крім того, чотири серед них (TMV, TSWV,
CMV, PVY) займають перші позиції в 10-ці найбільш нау-
ково і економічно значущих вірусів рослин в світі. Таке по-
ширення та шкодочинність цих вірусів пояснюється широ-
ким колом чутливих рослин-господарів, диких рослин і
бур'янів - резервантом інфекції, а також великою кількістю
векторів. Дослідження вірусологів з декількох країн пока-
зують, що симптоматика вірусних інфекцій ехінацеї з рока-
ми стає більш складною. Щорічно діагностуються нові ві-
руси ехінацеї, що ускладнює прогнозування та оцінку ризи-
ку появи епіфітотій в певних регіонах, як наприклад, це
було зареєстровано в Литві, Україні та Болгарії. У статті де-
тально представлені симптоми вірусних хвороб ехінацеї,
основні властивості кожного вірусу і дані про їхній вплив
на концентрацію біологічно активних речовин і важких ме-
талів рослин.
Ключов і с л ов а: ехінацея пурпурова, віруси рослин,
видове різноманіття, біологічно активні речовини, важкі
метали.
А. А. Дунич, Л. Т. Мищенко
Вирусы эхинацеи пурпурной:
видовое разнообразие и вредоносность
В мире все более острой проблемой при промышленном вы-
ращивании лекарственных растений становятся вирусные
заболевания. Установлено, что во многих странах число ви-
русов, поражающих эхинацею в последние годы значительно
возросло. Несмотря на это, цельный обзор мировой научной
литературы о вирусных заболеваниях, инфицирующих эту
ценную лекарственную культуру, отсутствует. Целью работы
является суммировать основную информацию о вирусах, за-
ражающих эхинацею пурпурную в мире. Анализ литературы
показал, что эхинацея пурпурная поражается 10 вирусами из
семейств Bromoviridae, Bunyaviridae, Secoviridae, Potyviridae,
Virgaviridae. Практически все они определены вредоносны-
ми фитовирусами. Кроме этого, четыре среди них (TMV,
TSWV, CMV, PVY) занимают первые позиции в 10-ке наибо-
лее научно и экономически значимых вирусов растений в
мире. Такое распространение и вредоносность этих вирусов
объясняется широким кругом чувствительных растений-хо-
зяев, диких растений и сорняков – резервантов инфекции, а
также большим количеством векторов. Исследования вирусо-
логов из нескольких стран показывают, что симптоматика
вирусных инфекций эхинацеи с годами становится более
сложной. Ежегодно диагностируются новые вирусы эхина-
цеи, что усложняет прогнозирование и оценку риска появле-
ния эпифитотий в определенных регионах, как например, это
было зарегистрировано в Литве, Украине и Болгарии. В ста-
тье детально представлены симптомы вирусных болезней
эхинацеи, основные свойства каждого вируса и данные о их
влиянии на концентрацию биологически активных веществ и
тяжелых металлов растениях.
Ключевые слова: эхинацея пурпурная, вирусы расте-
ний, видовое разнообразие, биологически активные вещес-
тва, тяжелые металлы.
Received 10.12.2014
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