Фенологічні зміни вториних метаболітів та мінерального живлення Solidago canadensis та їх вплив на ґрунтову екосистему ризосфери
A comprehensive study of the impact of Solidago canadensis on the soil ecosystem using the example of monodominant communities of this species located on the exposition plot ‘Steppes of Ukraine’ of the M.M. Gryshko National Botanical Garden of the NAS of Ukraine (Kyiv) is presented. The research was...
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M.M. Gryshko National Botanical Garden of the NAS of Ukraine
2025
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Plant Introduction| _version_ | 1860145171057868800 |
|---|---|
| author | Zaimenko, Natalia Chernikova, Nina Didyk, Nataliya Yunosheva, Olena Pavliuchenko, Nataliia Kharitonova, Iryna Malashchuk, Olena Zakrasov, Olexandr |
| author_facet | Zaimenko, Natalia Chernikova, Nina Didyk, Nataliya Yunosheva, Olena Pavliuchenko, Nataliia Kharitonova, Iryna Malashchuk, Olena Zakrasov, Olexandr |
| author_sort | Zaimenko, Natalia |
| baseUrl_str | https://www.plantintroduction.org/index.php/pi/oai |
| collection | OJS |
| datestamp_date | 2025-12-27T15:10:16Z |
| description | A comprehensive study of the impact of Solidago canadensis on the soil ecosystem using the example of monodominant communities of this species located on the exposition plot ‘Steppes of Ukraine’ of the M.M. Gryshko National Botanical Garden of the NAS of Ukraine (Kyiv) is presented. The research was conducted during the growing seasons of 2023–2024. The content of macro- and microelements, secondary metabolites in leaves and soil, allelopathic activity, organic and mineral carbon content, and laccase activity were assessed, and the functional structure of soil microbial communities was determined.It has been established that S. canadensis actively regulates the absorption of nutrients, which serves as an adaptive mechanism to environmental changes. The capacity of S. canadensis to reduce soil contamination with toxic metals such as Mn, Zn, Co, Cr, Fe, Pb, and Ti has been shown. For Mn and Zn, S. canadensis acts as a phytoextractor, and for Co, Cr, Fe, Pb, and Ti, it acts as a phytostabilizer.Significant phenological fluctuations in allelopathic activity, the content of biologically active secondary metabolites, and the functional structure of the rhizosphere microbiota in S. canadensis have been shown. The successful spread of S. canadensis outside its natural range may be associated with the ability of this plant to produce and release into the environment a number of secondary metabolites of phenols, terpenoids, saponins, etc., which are responsible for its high resistance to abiotic and biotic stress factors, allelopathic activity, inhibit the development of harmful bacteria and stimulate the development of actinomycetes, which participate in the restoration of soil fertility and are antagonists of phytopathogens. |
| doi_str_mv | 10.46341/PI2025007 |
| first_indexed | 2025-09-17T09:33:46Z |
| format | Article |
| fulltext |
© The Authors. This content is provided under CC BY 4.0 license.
Plant Introduction, 107, 36–47 (2025)
RESEARCH ARTICLE
Phenological changes in secondary metabolites and mineral nutrition of
Solidago canadensis and their impact on the rhizosphere soil ecosystem
Natalia Zaіmenko, Nina Chernikova, Nataliya Didyk *, Olena Yunosheva, Nataliia Pavliuchenko,
Iryna Kharytonova, Olena Malashchuk, Oleksandr Zakrasov
M.M. Gryshko National Botanical Garden, National Academy of Sciences of Ukraine, Sadovo-Botanichna str. 1, 01103 Kyiv, Ukraine;
* nataliya_didyk@ukr.net
Received: 30.04.2025 | Accepted: 27.08.2025 | Published online: 08.09.2025
Abstract
A comprehensive study of the impact of Solidago canadensis on the soil ecosystem using the example
of monodominant communities of this species located on the exposition plot ‘Steppes of Ukraine’ of
the M.M. Gryshko National Botanical Garden of the NAS of Ukraine (Kyiv) is presented. The research
was conducted during the growing seasons of 2023–2024. The content of macro- and microelements,
secondary metabolites in leaves and soil, allelopathic activity, organic and mineral carbon content, and
laccase activity were assessed, and the functional structure of soil microbial communities was determined.
It has been established that S. canadensis actively regulates the absorption of nutrients, which serves as an
adaptive mechanism to environmental changes. The capacity of S. canadensis to reduce soil contamination
with toxic metals such as Mn, Zn, Co, Cr, Fe, Pb, and Ti has been shown. For Mn and Zn, S. canadensis acts
as a phytoextractor, and for Co, Cr, Fe, Pb, and Ti, it acts as a phytostabilizer.
Significant phenological fluctuations in allelopathic activity, the content of biologically active secondary
metabolites, and the functional structure of the rhizosphere microbiota in S. canadensis have been shown.
The successful spread of S. canadensis outside its natural range may be associated with the ability of
this plant to produce and release into the environment a number of secondary metabolites of phenols,
terpenoids, saponins, etc., which are responsible for its high resistance to abiotic and biotic stress factors,
allelopathic activity, inhibit the development of harmful bacteria and stimulate the development of
actinomycetes, which participate in the restoration of soil fertility and are antagonists of phytopathogens.
Keywords: Solidago canadensis, mineral nutrition, allelopathic activity, secondary metabolites, rhizosphere soil, microbial communities,
phytoremediation
https://doi.org/10.46341/PI2025007
UDC [58.063.7 : 582.998.16] : 631.41 : 712.253 : 58.069.029(477-25)
Authors’ contributions: Zaimenko N. conceived and designed the experiments. Kharytonova I. and Malashchuk O. performed
agrochemical analysis of soil. Yunosheva O. performed microbiological analysis. Pavliuchenko N. performed allelopathic analysis of
soil. Chernikova N. and Didyk N. performed biochemical analysis of soil and plant material. Zasrasov O. determined laccase activity.
Chernikova N. and Didyk N. wrote the manuscript draft. Zaimenko N. critically revised the manuscript.
Funding: Departmental research program «The role of secondary metabolism products in the formation of allelopathic potential in
the soil - plant - soil system» 2023-2027, state registration number 0123U101379.
Competing Interests: The authors declare no conflict of interest.
ISSN 1605-6574, e-ISSN 2663-290X
https://creativecommons.org/licenses/by/4.0/
https://orcid.org/0000-0003-2379-1223
https://orcid.org/0009-0007-0188-4302
https://orcid.org/0000-0001-8448-7490
https://orcid.org/0000-0003-1349-7733
https://orcid.org/0000-0001-8934-7163
https://orcid.org/0000-0001-9540-5278
https://orcid.org/0000-0001-9762-8758
https://orcid.org/0009-0005-0753-5547
Plant Introduction • 107 37
Phenological changes in secondary metabolites and mineral nutrition of Solidago canadensis
Introduction
Climate change and urbanization promote the
spread of invasive species of plants and animals
altering biodiversity and the functioning of
natural and seminatural ecosystems. According
to EU Regulation 1143/2014 (EU 2014), invasive
species are alien species of plants or animals
whose introduction or spread threatens or
negatively affects biodiversity and ecosystem
services (Skočajić & Nešić, 2020).
Goldenrod (Solidago canadensis L.) is
classified as an invasive plant, which was
introduced to Europe as an ornamental from
North America in the early 17th century
(Radusiene et al., 2015). Since that time,
S. canadensis has spread rapidly in local
seminatural and natural ecosystems, which
resulted in its tremendous expansion and
displacement of native flora, especially
in various disturbed environments,
predominantly along roadsides and railways,
urban settings, abandoned fields, and
grasslands, forest edges, roadsides, and
meadows. Presently, it is considered one of
the most aggressive invaders, and is defined
by the European and Mediterranean Plant
Protection Organization as an invasive species
having a high potential for spread and posing
an important threat to the environment and
biodiversity in the region (EPPO) (Radusiene
et al., 2015). The EPPO (2024) recommends
taking control measures and raising awareness
due to S. canadensis high potential to alter
natural ecosystems and shift ecosystem
services.
Some authors indicated that S. canadensis
provides several benefits to its new habitats,
such as attraction of pollinators, forage
production, and storing carbon (Gallardo et al.,
2019). Besides, S. canadensis was shown to
stabilize increased (compared with natural)
concentrations of Pb and Zn in the soil
(Bielecka & Królak, 2019; Bielecka et al., 2020).
At the same time, it also accumulated Zn in the
above-ground parts, which allowed authors
(Bielecka & Królak, 2019; Bielecka et al., 2020)
to assume the possibility of using this plant
for phytoextraction of soils contaminated
with this metal. In addition, phytosanitary
properties of S .canadensis root secretions
were demonstrated, suppressing the growth
of phytopathogenic microorganisms such as
Botrytis cinerea (Liu et al., 2016), Fusarium sp.,
Phytophtora infestans, etc. (Anžlovar & Koce,
2014). Taking into account such traits of
S. canadensis as a wide range of tolerance to
soil physicochemical conditions and climatic
factors, ability to colonize contaminated soils,
and production of high biomass of above-
ground parts, an extensive underground
system makes this species a promising plant
for utilization for phytoremediation and
phytosanation of contaminated soils with toxic
metals/phytopatogens (Bielecka & Królak,
2019).
The extraction of new products from the
invasive species presents a viable approach
aimed at their valorisation. In recent years,
significant attention has been focused on
phytochemicals of Solidago species, including
S. canadensis (Poljuha et al., 2024). The
latter was shown to contain a diverse array
of bioactive compounds with antioxidant,
antimicrobial, anticancer, anti-inflammatory,
and hepatoprotective activity. Phytоchemicals
of S. canadensis can be used in pharmacy,
agriculture, medicine, cosmetics, production
of natural dyes, cellulose, biopesticides, etc.
(Poljuha et al., 2024).
Solidago canadensis exhibits strong
allelopathic effects, inhibiting the growth
and development of native species, thereby
reducing biodiversity. The allelochemicals
responsible for the described effects identified
are chlorogenic acid, rutin (quercetin-3-O-
rutinoside), kaempferol-3-O-D-glucoside, and
quercitrine released through root exudates,
plant residue decomposition, and leachates,
which alter soil microbial communities and
inhibit arbuscular mycorrhizal fungi, vital for
nutrient and water uptake in native plants
(Likhanov et al., 2021; Kato-Noguchi & Kato,
2022; Zhu et al., 2022). These exometabolites
enhance species invasiveness and present
promise as potential natural herbicides (Yang
& Li, 2022).
Many scientific papers are devoted to
the impact of S. canadensis on biodiversity
decline (Wang et al., 2018a, 2018b, 2019, 2021;
Gallardo et al., 2019). While there is a lack of
comprehensive information on S. canadensis
effect on soil chemical and physical properties,
biochemical and allelopathic regime as well as
on how S. canadensis allelochemicals alter soil
microbial communities.
This study aimed to conduct a complex
assessment of S. canadensis effects on the
38 Plant Introduction • 107
Zaіmenko et al.
soil ecosystem, including metal cations,
biologically active secondary metabolites, soil
microbial community, allelopathic regime, and
organic carbon deposition.
Material and methods
The research objects are rhizosphere soil
and S. canadensis plants growing in the
‘Steppes of Ukraine’ exposition plot of the
M.M. Gryshko National Botanical Garden
of the NAS of Ukraine (Kyiv). Sampling was
made during the period of S. canadensis shoot
development (early June), flowering (end of
July), and the seed maturation stage (middle of
September) in 2023–2024. As the control, we
used the soil collected from the rhizosphere
of the surrounding forb meadow communities
dominated by indigenous species Elytrigia
intermedia (Host) Nevski, Adonis vernalis L.,
and Paeonia tenuifolia L. A comparative
analysis of the content of chemical elements
in the soil and in the above-ground parts
of plants was carried out, the functional
structure of soil microbial communities was
determined, and the allelopathic activity of the
above-ground parts and rhizosphere soil was
assessed. Soil samples were taken with a drill
at 10–20 cm depth, sieved through a sieve with
openings of 1–2 mm.
Agronomic soil analysis
To determine the pH of the soil solution, a
portion of the sieved and dry soil was placed
in a porcelain beaker and poured with 25 ml
of 1 N KCl (pH 5.5–6.0) and kept for 12 h, after
which measurements were performed using
the MW 402 TDS Meter (Milwaukee, Hungary).
The preparation of the soil samples for
the analysis of the content of macro- and
microelements was carried out according to
Zaimenko et al. (2021). Acid-soluble forms of
metal cations and other chemical elements
were extracted using a 1.0 N solution of
nitric acid (HNO3). Preparation of plant
samples for analysis was done by wet ashing
with a solution of high-purity nitric acid
using Speedwave Xpert DAP-60X (Berghof,
Germany), a special system for microwave
decomposition of samples. The concentrations
of chemical elements in the solution were
measured using an ICAP 6300 DUO plasma
emission spectrometer (Thermo Fisher
Scientific, USA). The relative uncertainty
ranged from 7 % to 12 % for different chemical
elements. Laccase activity was determined
spectrophotometrically using the color
reaction with syringaldazine according to
Dzyuba et al. (2021) using a SPECORD 200
instrument (Analytik Jena, Germany).
Allelopathic analysis
Water-soluble allelochemicals from freshly
harvested crushed leaves of S. canadensis were
extracted with distilled water (1 : 100) for 4
hours. This concentration corresponds to the
observed levels of goldenrod allelochemicals
in its monodominant communities in the field
(Yuan et al., 2013). The allelopathic activity
of the extracts was assessed by bioassay on
radish (Raphanus sativus L. var. sativus) root
growth (Pavliuchenko et al., 2021; Pavliuchenko
& Young, 2021). The allelopathic activity of
S. canadensis rhizosphere soil was assessed
by direct bioassay using watercress (Lepidium
sativum L.) as a test plant (Pavliuchenko
et al. 2021; Pavliuchenko & Young, 2021). The
percentage of root growth inhibition was
measured by the given formula:
GI = 100×(PC–PT)/PC, where
GI – growth inhibition, in %,
PT – length of roots of seedlings grown with
extracts (treatment group), in cm,
PC – length of the roots of seedlings in the
control group, in cm.
Microbiological analysis
The functional structure of microbial
communities was assessed by seeding soil
samples on selective media. In particular,
micromycetes were determined on Čapek’s
medium, actinomycetes on starch-ammonia
agar (KAA), ammonifiers on meat-peptone
agar (MPA), and microorganisms that consume
mainly mineral nitrogen compounds on KAA.
The growth of nitrogen-fixing microorganisms
was taken into account by the percentage
of soil clods fouling on the Ashby medium
(Ellanska et al., 2021).
The number of microorganisms was
determined by direct colony counting in Petri
dishes. The results were expressed in colony-
forming units (CFU) per 1 g of soil. A sample
dilution was analyzed to obtain reliable data,
Plant Introduction • 107 39
Phenological changes in secondary metabolites and mineral nutrition of Solidago canadensis
in which 10 to 150 colonies were formed on a
nutrient medium in a Petri dish (Ellanska et al.,
2021).
The number of microorganisms was
calculated per 1 g of dry soil. To do this, the
moisture content was first determined: a
sample weighing 10 g was placed in a pre-
weighed metal box and dried in a drying oven
at 105 °C for 3 hours.
The number of microorganisms was
calculated using the formula:
а =b/(c × d × f), where
a – the number of cells in 1 g of dry soil;
b – the average number of colonies from
one Petri dish;
c – the dilution from which the culture was
made, in ml;
d – the volume of suspension that was
applied to the nutrient medium, in ml;
f – the mass of soil that was taken for
analysis, in g.
Biochemical analysis of plant material and soil
Low molecular weight phenolic compounds
were extracted from plant material first
with distilled water and then with 80 %
ethanol. The content was determined
spectrophotometrically using a color reaction
with Folin-Ciocalteu reagent (Pavliuchenko
et al., 2021; Pavliuchenko & Young, 2021).
Flavonoids were extracted with 70 % ethanol;
the quantitative content was determined
spectrophotometrically using a color reaction
with 10 % AlCl3 solution (Pavliuchenko et al.,
2021; Pavliuchenko & Young, 2021). Tannins
were extracted with boiling distilled water.
The quantitative content was determined by
titration with 0.1 % potassium permanganate
solution in the presence of indigocarmine
(Mardar & Serdyuk, 2008). Triterpenoids and
saponins were extracted with 96 % ethanol,
and the quantitative content was determined
spectrophotometrically using a color reaction
with vanillin reagent (Pavliuchenko et al., 2021;
Pavliuchenko & Young, 2021).
The free phenolic compounds are isolated
from the soil by ion exchange (desorption)
with the use of ion exchanger KU-2-8
(H+) as a model of the root system with
dissolving and absorbing ability in relation
to mobile organic compounds (Pavliuchenko
et al., 2021; Pavliuchenko & Young, 2021). All
spectrophotometric measurements were
performed using the instrument SPECORD
200 (Analytik Jena, Germany).
Statistical analysis
The results are presented as mean ± standard
error (m ± SE). The reliability of the difference
(P < 0.05) between the obtained data was
determined by the method of variance analysis
(one-factor variance analysis) using Tukey’s a
posteriori test using MS Excel and Statistica
10.0 software (Stat-Soft Inc., USA) for the data
processing.
Results and discussion
Comparative analysis of the content of
chemical elements in soil and plants
S. canadensis indicated good ability of this
species to regulate exchange of macro- and
microelements with the soil environment
(Tables 1 & 2). Of practical interest is the ability
of S. canadensis to significantly reduce the
level of such toxic metals as Co, Cr, Fe, Mn,
Pb, Ti, and Zn in the soil. While the content
of Al and P in the rhizosphere of S. canadensis
increased 3.8 and 2 times from the beginning
to the end of the vegetation season.
The phenological dynamics of the
content of the metal cations in the leaves of
S. canadensis are mainly due to phenological
changes in physiological processes
associated with active vegetative growth,
flowering, and seed formation, as well as
weather conditions (high temperature and
drought in mid-summer and early autumn).
In particular, the concentration of Ca, which
is known to enhance plant tolerance towards
abiotic stress (Gupta et al., 2023), reached
maximum values in September and was 3.1
times higher compared to the initial phase
of plant development. The concentration of
P, essential for flower and seed development,
was minimal during the vegetative phase and
reached a maximum at full flowering. The
presence of vanadium in the leaves in the
absence of detectable concentrations of this
element in the soil.
The decrease in Mn and Zn concentrations
in the soil correlated with the increase in
the accumulation of these toxic metals in
the above-ground parts of S. canadensis,
which indicates the possibility of using this
40 Plant Introduction • 107
Zaіmenko et al.
invasive species for phytoextraction of these
pollutants from the soil. Also, S. canadensis
is promising for phytostabilization of toxic
metals, such as Co, Cr, Fe, Pb, and Ti, the
concentrations of which were reduced in
the soil but did not increase in the above-
ground parts. The potential of S. canadensis
to adsorb Pb and Zn from the surrounding
soil was shown by Bielecka & Królak (2019).
The authors demonstrated the good ability of
this plant to actively transport metals through
the membranes of root cells, chelate them
using organic acids, flavonoids, and phenolic
compounds, and store metal chelates in cell
vacuoles to minimize toxic effects. The high
resistance of S. canadensis to heavy metal
contamination (zinc and lead) was also shown
by Czortek et al. (2020).
Analysis of the allelopathic activity of
aqueous extracts from freshly collected
material and rhizosphere soil of S. canadensis
revealed the presence of inhibitors of growth
processes of acceptor plants both in the leaves
and in the roots of the donor plant (Fig. 1). The
phenological changes in the accumulation of
inhibitors in plants had a parabolic character
with a maximum in July (flowering phase) and a
minimum in September (seed ripening phase).
Meanwhile, the phenological dynamics of soil
allelopathic activity had a U-shaped pattern,
with a maximum during vegetative growth,
a sharp drop in the flowering phase, and a
subsequent rise in autumn. The allelopathic
activity of aqueous extracts from leaves and
rhizosphere soil of S. canadensis was positively
correlated with the content of low-molecular
phenolic substances (Table 3), which indicates
the important role of these substances in
the formation of the allelopathic potential of
S. canadensis.
Element
S. canadensis Meadow forbs
Shoot
development
Flowering Seed
maturation
Shoot
development
Flowering Seed
maturation
Al 1067.0 ± 53.1 7028.0 ± 315.3 4047.0 ± 196.3 11995.0 ± 132.8 13440.0 ± 41.5 17070.0 ± 148.2
B 9.5 ± 0.5 7.09 ± 0.4 1.4 ± 0.09 7.1 ± 0.6 7.2 ± 0.6 11.3 ± 1.1
Ba 76.6 ± 3.5 79.3 ± 3.5 35.5 ± 1.8 67.3 ± 2.9 43.3 ± 2.1 39.267.3 ± 2.4
Ca 3250.0 ± 152.6 2328.0 ± 113.5 2456.0 ± 120.6 3104.0 ± 273.5 3628.8 ± 161.8 3373.7 ± 221.8
Co 4.0 ± 0.2 3.4 ± 0.2 1.4 ± 0.1 5.6 ± 0.9 5.1 ± 0.4 4.2 ± 0.6
Cr 14.0 ± 0.7 11.8 ± 0.4 4.4 ± 0.2 24.1 ± 1.1 22.9 ± 1.2 38.3 ± 1.6
Cu 15.4 ± 0.7 15.5 ± 0.8 9.4 ± 0.4 15.8 ± 1.2 24.4 ± 2.1 21.5 ± 1.7
Fe 7086.0 ± 348.2 5763.0 ± 282.4 2967.0 ± 141.83 2563 ± 129.1 2465.8 ± 137.3 2257.4 ± 122.2
К 2379.0 ± 155.3 1239.0 ± 59.8 557.3 ± 26.3 116.4 ± 65.6 174.2 ± 77.1 128.4 ± 46.1
Mg 1492.0 ± 74.5 903.8 ± 44.3 516.8 ± 24.3 828.7 ± 39.5 1258.0 ± 54.1 378.8 ± 42.7
Mn 532.9 ± 25.2 271.0 ± 12.9 119.4 ± 5.9 503.2 ± 19.3 477.0 ± 28.5 416.6 ± 32.1
Na 64.8 ± 3.2 50.3 ± 2.5 32.4 ± 1.6 89.0 ± 2.6 59.8 ± 3.4 55.2 ± 2.1
Ni 8.2 ± 0.4 9.0 ± 0.4 4.4 ± 0.2 9.9 ± 0.5 7.1 ± 0.5 6.8 ± 0.4
P 3.7 ± 0.2 8.4 ± 0.4 7.4 ± 0.3 9.2 ± 0.1 15.3 ± 0.3 12.2 ± 0.1
S 703.0 ± 33.3 557.0 ± 24.8 887.8 ± 41.2 188.6 ± 28.6 143.4 ± 32.5 156.3 ± 20.1
Pb 74.6 ± 0.7 68.7 ± 3.3 34.5 ± 1.6 122.9 ± 2.7 118.5 ± 2.4 109.4 ± 3.2
Si 1002.0 ± 48.8 399.9 ± 18.9 132.5 ± 5.9 1438.0 ± 98.4 1062.0 ± 92.3 837.0 ± 33.6
Sr 16.7 ± 0.8 10.4 ± 0.5 14.5 ± 0.7 19.6 ± 0.8 16.9 ± 0.2 15.0 ± 0.4
Ti 331.0 ± 15.7 129.2 ± 6.0 15.5 ± 0.7 556.4 ± 12.2 495.0 ± 7.1 432.9 ± 5.3
Zn 41.4 ± 1.7 4.8 ± 2.0 21.3 ± 1.1 47.0 ± 2.3 30.0 ± 1.9 46.0 ± 2.5
Table 1. Phenological changes in the content of chemical elements in the rhizosphere soil of Solidago
canadensis and meadow forbs, mg/kg (averages for 2023–2024).
Plant Introduction • 107 41
Phenological changes in secondary metabolites and mineral nutrition of Solidago canadensis
Element
Phenological stage
Shoot
development
Flowering Seed
maturation
Al 295.7 ± 14.78 120.3 ± 5.87 215.8 ± 9.96
B 33.8 ± 1.45 42.9 ± 2.03 45.8 ± 2.14
Ba 6.01 ± 0.27 11.27 ± 0.51 9.63 ± 0.42
Ca 5705 ± 279.3 8799 ± 421.7 18080 ± 892.8
Co 0.34 ± 0.01 0.33 ± 0.01 0.34 ± 0.01
V 5.51 ± 0.24 6.08 ± 10.29 6.41 ± 0.31
Cr 1.65 ± 0.08 1.55 ± 0.08 1.40 ± 0.07
Cu 10.81 ± 0.52 9.62 ± 0.46 9.99 ± 0.48
Fe 256.2 ± 12.77 130.2 ± 6.43 209.9 ± 10.35
К 23910 ± 1147 12330 ± 587 9828 ± 493
Mg 1880 ± 95.2 1798 ± 87.4 2153 ± 103.6
Mn 69.74 ± 3.51 68.05 ± 3.42 80.24 ± 3.87
Na 37.70 ± 1.75 28.39 ± 1.42 55.77 ± 2.69
Ni 2.12 ± 0.11 2.18 ± 0.10 1.37 ± 0.06
P 20.58 ± 1.03 43.9 ± 2.11 34.7 ± 1.69
S 2300 ± 109.7 755 ± 35.9 1819 ± 89.7
Pb 2.47 ± 0.12 1.83 ± 0.08 1.10 ± 0.05
Si 1369 ± 67.5 441.6 ± 22.1 733.1 ± 35.9
Sr 6.16 ± 0.29 15.75 ± 0.76 26.33 ± 1.28
Ti 11.93 ± 0.52 2.46 ± 0.12 6.76 ± 0.32
Zn 18.53 ± 0.88 37.38 ± 1.83 38.92 ± 0.91
Table 2. Phenological changes in the content of
chemical elements in Solidago canadensis leaves,
mg/kg (average for 2023–2024).
Strong inhibitory effect of S. canadensis
allelochemicals to native flora was shown in
many studies (Zhu et al., 2022; Kato-Noguchi
& Kato, 2022; Yang & Li, 2022). The allelopathic
effect of S. canadensis is associated with
phenolic acids (ferulic, p-coumaric, caffeic,
chlorogenic acid, etc.), flavonoids (kaempferol,
quercitrin, and rutin), fatty acids, terpenes,
polyphenols, and saponins (Likhanov et al.,
2021; Zhu et al., 2022; Kato-Noguchi & Kato,
2022). The results of our research confirm the
opinion of other authors that the allelopathy of
S. canadensis may contribute to its increasing
competitive ability and make the plant invasive.
Analysis of the secondary metabolites
of S. canadensis showed a high content of
biologically active compounds related to
phenols and terpenoids, indicating its powerful
adaptive potential and ecological plasticity.
Flavonoids are potent antioxidants that
neutralize reactive oxygen species (ROS) and
protect cells from oxidative stress (Mierziak
et al., 2014). The high content of flavonoids in
the leaves of S. canadensis indicates its ability
to effectively counteract abiotic stress factors,
such as drought, high temperatures, or UV
radiation. Some flavonoids exhibit allelopathic
effects, inhibiting the growth of other plants,
which may explain the invasive ability of
goldenrod in phytocenoses (Mierziak et al.,
2014).
Tannins can bind proteins, making the plant
less attractive to herbivores and insect pests
Figure 1. Phenological changes in allelopathic activity of 1 % aqueous extracts from leaves and roots of
Solidago canadensis (bioassay for radish root growth) and rhizosphere soil (cress root growth), growth
inhibition in %. Phenological stages: I – shoot development, II – flowering, III – seed maturation.
42 Plant Introduction • 107
Zaіmenko et al.
(Barbehenn & Constabel, 2011). Their high
concentration in the leaves of S. canadensis
may be a key factor in the formation of
resistance to phytopathogens. The high
concentration of saponins and triterpenoids in
the leaves may also be a defense mechanism
against phytophagous organisms (Mathur
et al., 2025).
The antimicrobial activity of S. canadensis
is thought to be due to the terpenoids present
in the root essential oils and soluble phenolic
and polyphenolic compounds (Anžlovar
& Koce, 2014). The results of our studies
showed the presence of biologically active
concentrations of saponins, triterpenoids, and
especially phenolics in the rhizosphere soil of
S. canadensis throughout the growing season,
with maximums in July (for triterpenoids) and
September (for phenolics).
Solidago canadensis phytotoxic
allelochemicals not only suppress other plants’
growth, but also alter the community structure
of soil microbiota (Table 4).
Comparison of the results of the assessment
of the biochemical regime of the S. canadensis
rhizosphere soil with the changes in the
functional structure of microbial communities
showed that a 2.9-fold increase in the content
of low-molecular-weight phenolics in the
rhizosphere soil was accompanied by a 1.7-fold
decrease in the number of ammonifiers, and an
increase in the number of actinomycetes and
micromycetes by 2 and 1.14 times, respectively.
This trend is consistent with the findings of
other authors that phenolic allelochemicals
inhibit the development of soil bacteria but
stimulate the growth of microfungi and
actinomycetes (Zhou et al., 2012; Badri et al.,
2013).
Actinomycetes are also known for their
ability to produce and release into the
environment phenolic secondary metabolites
with diverse biological activity (Golińska &
Dahm, 2011). Therefore, the autumn increase in
phenolic content in S. canadensis rhizosphere
soil could result from the intensification of
actinomycetes metabolic activity, among which
many antibiotic compounds are produced.
Today, actinomycetes are recognized as
noteworthy antibiotic producers, making
40 % of the 160 microbial-based antibiotics
(Raut et al., 2023). Actinomycetes are also an
important component of the soil ecosystem
because of their ability to decompose many
complex compounds, such as proteins,
pectins, cellulose, hemicellulose, lignins, and
chitin (Golińska & Dahm, 2011).
The ecological outcome of microbiological
Secondary metabolites
Phenological stage
Shoot development Flowering Seed maturation
Leaves
Total phenolics 41.3 ± 3.4 77.4 ± 3.3 54.2 ± 2.7
Flavonoids 22.6 ± 2.8 56.6 ± 3.4 42.3 ± 2.9
Tannins 49.8 ± 2.9 76.2 ± 3.4 111.3 ± 3.5
Saponins 4.7 ± 0.2 12.2 ± 0.7 9.5 ± 0.9
Triterpenoids 6.5 ± 0.6 13.8 ± 0.8 17.6 ± 1.1
Rhizosphere soil
Triterpenoids 1.9 ± 0.2 2.6 ± 0.1 1.8 ± 0.1
Saponins 1.2 ± 0.1 1.8 ± 0.2 1.7 ± 0.1
Total phenolics 58.73 ± 1.9 24.8 ± 2.2 71.9 ± 4.8
Control soil
Triterpenoids 1.6 ± 0.1 2.1 ± 0.1 1.4 ± 0.1
Saponins 1.4 ± 0.2 1.6 ± 0.1 1.3 ± 0.1
Total phenolics 51.4 ± 3.5 67.2 ± 3.1 78.6 ± 2.2
Table 3. Content of secondary metabolites in leaves and the rhizosphere soil of Solidago canadensis,
mg/g DW.
Plant Introduction • 107 43
Phenological changes in secondary metabolites and mineral nutrition of Solidago canadensis
Plants Phenological
stage
Micromycetes,
thousands
CFU per g DW
soil
Actinomycetes,
millions CFU
per g DW soil
Ammonifiers,
millions CFU
per g DW soil
Microorganisms
that consume
mineral
nitrogen,
millions CFU
per g DW soil
Mineralization
coefficient
S. canadensis Flowering 34.7 ± 2.9 0.7 ± 0.07 6.1 ± 0.2 6.3 ± 0.1 1.0
Seed
maturation
39.7 ± 1.5 1.4 ± 0.2 3.5 ± 0.1 6.1 ± 0.1 1.7
Meadow
forbs
Flowering 58.4 ± 1.1 1.34 ± 0.1 7.8 ± 0.1 8.8 ± 0.2 1.4
Seed
maturation
43.9 ± 1.3 1.15 ± 0.1 4.6 ± 0.2 9.4 ± 0.1 1.8
Table 4. The number of microorganisms belonging to different functional groups in the rhizosphere soil
of Solidago canadensis and meadow forbs.
processes was assessed by the ratio of the
number of microorganisms with mineral
and organic nutrition. The maximum
mineralization-immobilization coefficient
in the S. canadensis rhizosphere soil was
observed at the end of the growing season.
This tendency was confirmed by the results of
the analysis of the content of organic carbon
and laccase activity (Table 5).
The significant impact of S. canadensis on
soil microflora has been shown in the works of
other authors. In particular, Qiao et al. (2024)
found that the S. canadensis enhanced nutrient-
releasing microorganisms (Actinomycetota)
and disease-resistant microorganisms
(Nocardioides), while decreasing N-cycling
microorganisms (Nitrososphaeria and
Nitrospirota). These changes in the soil
microbiome create conditions that promote
the effective use of S. canadensis resources
and its aggressive invasion. The ability of
S. canadensis to influence nitrogen-fixing
bacterial communities and adapt to varying
levels of heavy metals in the soil may also
contribute to its invasive success (Wang et al.,
2018a, 2018b, 2023).
It is known that metabolites of S. canadensis
can inhibit the activity of microorganisms
responsible for the nitrogen cycle and promote
the dominance of microorganisms that are
adapted to allelopathic substances (Zhu et al.,
2022; Kato-Noguchi & Kato, 2022; Wang et al.,
2023). Also, allelochemicals of S. canadensis
are known for their toxicity to insect pests,
affecting their nervous system, reproductive
ability, and development (Benelli et al., 2019).
The content of soil organic carbon is an
indicator of its fertility, health, and ability to
provide ecosystem services. According to the
results of our research, the content of organic
carbon in the rhizosphere soil of S. canadensis
is quite high throughout the growing season,
if compared with the organic carbon level
common for the meadow-steppe vegetation
in our region (Vyshenska & Ivanyk, 2015;
Zaimenko et al., 2022), which confirms the
positive role of this plant in the deposition of
atmospheric carbon dioxide in the form of soil
organic carbon.
Laccases are oxidoreductase enzymes
with polyphenol oxidase activity that belong
to the multicopper oxidase superfamily (Aza
& Camarero, 2023). After being discovered
in the exudates of the oriental lacquer
tree, Toxicodendron vernicifluum (Stokes)
F.A. Barkley, laccases have been identified in
fungi, bacteria, and insects (Aza & Camarero,
2023). Presently, laccases are considered
promising for the remediation of water and
soil environment due to their ecological safety
and ability to oxidize a wide range of aromatic
pollutants to less toxic derivatives (Aza &
Camarero, 2023). The relatively high laccase
activity in the rhizosphere of S. canadensis
throughout the growing season explains the
resistance of this plant to its own phenolic
allelochemicals and other organic aromatic
pollutants. It should be noted that Solidago
plants were shown to be capable of absorbing,
translocating, and accumulating in foliar tissues
organic pollutants involved in the manufacturing
of explosives (Groom et al., 2002).
Note. CFU – colony-forming unit.
44 Plant Introduction • 107
Zaіmenko et al.
Conclusions
The effective spread of S. canadensis outside
its natural range is associated with the
ability of this plant to produce and release
into the environment a number of secondary
metabolites such as phenols, terpenoids,
saponins, etc., which are responsible for its
high resistance to abiotic and biotic stress
factors, create favorable soil conditions for
this species – reduce soil pollution with
toxic metals (Mn, Zn, Со, Сr, Fe, Pb, and
Ti), inhibit the development of other plant
species, harmful bacteria and stimulate the
development of actinomycetes, which take
part in the restoration of soil fertility and
are antagonists of phytopathogens. Analysis
of literature data, as well as the results of
our research, allows us to conclude that the
use of monodominant stands of this species
for phytoremediation and restoration of
fertility of soils contaminated with toxic
metals, aromatic organic and inorganic
toxicants, bacterial phytopathogens, and
insect pests is promising. Due to its high
productivity and resistance, S. canadensis
exerts a powerful influence on the
chemical, biochemical, allelopathic, and
microbiological characteristics of the
soil throughout the growing season. The
impact of soil conditions, microbiota, and
specific types of anthropogenic load on
the effectiveness of the phytoremediation
potential of S. canadensis, as well as the
possibility of using local flora species to
control the unwanted spread of this invasive
species, require further study.
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Фенологічні зміни вториних метаболітів та мінерального живлення Solidago
canadensis та їх вплив на ґрунтову екосистему ризосфери
Наталія Заіменко, Ніна Чернікова, Наталія Дідик *, Олена Юношева, Наталія Павлюченко, Ірина
Харитонова, Олена Малащук, Олександр Закрасов
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Plant Introduction • 107 47
Phenological changes in secondary metabolites and mineral nutrition of Solidago canadensis
активність лакази, а також з’ясували функціональну структуру мікробних угруповань ґрунту.
Встановлено, що S. canadensis активно регулює поглинання поживних елементів, що слугує
адаптивним механізмом до змін довкілля. Показано здатність S. canadensis знижувати забруднення
ґрунту токсичними такими металами як Mn, Zn, Со, Сr, Fe, Pb, Ti. При цьому для Mn та Zn S. canadensis
виступає у ролі фіто екстрактора, а для Со, Сr, Fe, Pb, Ti – фітостабілізатора.
Показано значні фенологічні коливання алелопатичної активності, вмісту біологічно активних
вторинних метаболітів та функціональної структури мікробіоти ризосфери у S. canadensis. Успішне
розповсюдження S. canadensis за межами його природного ареалу може бути пов`язане зі здатністю
цієї рослини продукувати та виділяти у навколишнє середовище низку вторинних метаболітів
фенольних, терпеноїдів сапонінів та ін., які відповідають за його високу стійкість до абіотичних і
біотичних стресових чинників, алелопатичну активність, пригнічують розвиток шкідливих бактерій
та стимулюють розвиток актиміцетів, які приймають участь у відновленні родючості ґрунту та є
антагоністами фітопатогенів.
Ключові слова: Solidago canadensis, мінеральне живлення, алелопатична активність, вторинні метаболіти, ризосферний
ґрунт, мікробні угруповання, фіторемедіація
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| id | oai:ojs2.plantintroduction.org:article-1658 |
| institution | Plant Introduction |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2026-02-08T08:11:56Z |
| publishDate | 2025 |
| publisher | M.M. Gryshko National Botanical Garden of the NAS of Ukraine |
| record_format | ojs |
| resource_txt_mv | wwwplantintroductionorg/32/4eee1265e381c0594f1c88846f4f6e32.pdf |
| spelling | oai:ojs2.plantintroduction.org:article-16582025-12-27T15:10:16Z Phenological changes in secondary metabolites and mineral nutrition of Solidago canadensis and their impact on the rhizosphere soil ecosystem Фенологічні зміни вториних метаболітів та мінерального живлення Solidago canadensis та їх вплив на ґрунтову екосистему ризосфери Zaimenko, Natalia Chernikova, Nina Didyk, Nataliya Yunosheva, Olena Pavliuchenko, Nataliia Kharitonova, Iryna Malashchuk, Olena Zakrasov, Olexandr A comprehensive study of the impact of Solidago canadensis on the soil ecosystem using the example of monodominant communities of this species located on the exposition plot ‘Steppes of Ukraine’ of the M.M. Gryshko National Botanical Garden of the NAS of Ukraine (Kyiv) is presented. The research was conducted during the growing seasons of 2023–2024. The content of macro- and microelements, secondary metabolites in leaves and soil, allelopathic activity, organic and mineral carbon content, and laccase activity were assessed, and the functional structure of soil microbial communities was determined.It has been established that S. canadensis actively regulates the absorption of nutrients, which serves as an adaptive mechanism to environmental changes. The capacity of S. canadensis to reduce soil contamination with toxic metals such as Mn, Zn, Co, Cr, Fe, Pb, and Ti has been shown. For Mn and Zn, S. canadensis acts as a phytoextractor, and for Co, Cr, Fe, Pb, and Ti, it acts as a phytostabilizer.Significant phenological fluctuations in allelopathic activity, the content of biologically active secondary metabolites, and the functional structure of the rhizosphere microbiota in S. canadensis have been shown. The successful spread of S. canadensis outside its natural range may be associated with the ability of this plant to produce and release into the environment a number of secondary metabolites of phenols, terpenoids, saponins, etc., which are responsible for its high resistance to abiotic and biotic stress factors, allelopathic activity, inhibit the development of harmful bacteria and stimulate the development of actinomycetes, which participate in the restoration of soil fertility and are antagonists of phytopathogens. Представлено комплексне дослідження впливу життєдіяльності рослин Solidago canadensis на ґрунтову екосистему на прикладі монодомінантних угруповань цього виду на ділянці “Степи України” Національного ботанічного саду ім. М.М. Гришка НАН України (м. Київ). Дослідження проводили впродовж вегетаційних сезонів 2023–2024 рр. Визначили вміст макро- і мікроелементів, вторинних метаболітів у листках та ґрунті, алелопатичну активність, вміст органічного та мінерального вуглецю, активність лакази, а також з’ясували функціональну структуру мікробних угруповань ґрунту.Встановлено, що S. canadensis активно регулює поглинання поживних елементів, що слугує адаптивним механізмом до змін довкілля. Показано здатність S. canadensis знижувати забруднення ґрунту токсичними такими металами як Mn, Zn, Со, Сr, Fe, Pb, Ti. При цьому для Mn та Zn S. canadensis виступає у ролі фіто екстрактора, а для Со, Сr, Fe, Pb, Ti – фітостабілізатора.Показано значні фенологічні коливання алелопатичної активності, вмісту біологічно активних вторинних метаболітів та функціональної структури мікробіоти ризосфери у S. canadensis. Успішне розповсюдження S. canadensis за межами його природного ареалу може бути пов`язане зі здатністю цієї рослини продукувати та виділяти у навколишнє середовище низку вторинних метаболітів фенольних, терпеноїдів сапонінів та ін., які відповідають за його високу стійкість до абіотичних і біотичних стресових чинників, алелопатичну активність, пригнічують розвиток шкідливих бактерій та стимулюють розвиток актиміцетів, які приймають участь у відновленні родючості ґрунту та є антагоністами фітопатогенів. M.M. Gryshko National Botanical Garden of the NAS of Ukraine 2025-09-08 Article Article application/pdf https://www.plantintroduction.org/index.php/pi/article/view/1658 10.46341/PI2025007 Plant Introduction; No 107 (2025); 36-47 Інтродукція Рослин; № 107 (2025); 36-47 2663-290X 1605-6574 en https://www.plantintroduction.org/index.php/pi/article/view/1658/1573 Copyright (c) 2025 Natalia Zaimenko, Nina Chernikova, Nataliya Didyk, Olena Yunosheva, Nataliia Pavliuchenko, Iryna Kharitonova, Olena Malashchuk, Olexandr Zakrasov http://creativecommons.org/licenses/by/4.0 |
| spellingShingle | Zaimenko, Natalia Chernikova, Nina Didyk, Nataliya Yunosheva, Olena Pavliuchenko, Nataliia Kharitonova, Iryna Malashchuk, Olena Zakrasov, Olexandr Фенологічні зміни вториних метаболітів та мінерального живлення Solidago canadensis та їх вплив на ґрунтову екосистему ризосфери |
| title | Фенологічні зміни вториних метаболітів та мінерального живлення Solidago canadensis та їх вплив на ґрунтову екосистему ризосфери |
| title_alt | Phenological changes in secondary metabolites and mineral nutrition of Solidago canadensis and their impact on the rhizosphere soil ecosystem |
| title_full | Фенологічні зміни вториних метаболітів та мінерального живлення Solidago canadensis та їх вплив на ґрунтову екосистему ризосфери |
| title_fullStr | Фенологічні зміни вториних метаболітів та мінерального живлення Solidago canadensis та їх вплив на ґрунтову екосистему ризосфери |
| title_full_unstemmed | Фенологічні зміни вториних метаболітів та мінерального живлення Solidago canadensis та їх вплив на ґрунтову екосистему ризосфери |
| title_short | Фенологічні зміни вториних метаболітів та мінерального живлення Solidago canadensis та їх вплив на ґрунтову екосистему ризосфери |
| title_sort | фенологічні зміни вториних метаболітів та мінерального живлення solidago canadensis та їх вплив на ґрунтову екосистему ризосфери |
| url | https://www.plantintroduction.org/index.php/pi/article/view/1658 |
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