Діагностичне значення синтезу фенольних сполук і проліну в листках Schisandra chinensis та Actinidia arguta для індикації рівня стресу рослин в умовах змішаних насаджень
The peculiarities of the accumulation of polyphenolic compounds and free proline were investigated in the leaves of Actinidia arguta and Shisandra chinensis during their cultivation in vegetation containers with different ratios of the number of plants, namely 1 : 1, 2 : 1, 1 : 2. Monocultural (sing...
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| Дата: | 2022 |
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M.M. Gryshko National Botanical Garden of the NAS of Ukraine
2022
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Plant Introduction| _version_ | 1860145116880044032 |
|---|---|
| author | Venediktova, Tetyana Zaimenko, Natalia Skrypchenko, Nadiia |
| author_facet | Venediktova, Tetyana Zaimenko, Natalia Skrypchenko, Nadiia |
| author_sort | Venediktova, Tetyana |
| baseUrl_str | https://www.plantintroduction.org/index.php/pi/oai |
| collection | OJS |
| datestamp_date | 2023-08-26T20:38:56Z |
| description | The peculiarities of the accumulation of polyphenolic compounds and free proline were investigated in the leaves of Actinidia arguta and Shisandra chinensis during their cultivation in vegetation containers with different ratios of the number of plants, namely 1 : 1, 2 : 1, 1 : 2. Monocultural (single-species) planting was used as a control. The content of free proline in plant leaves was carried out according to the method, which is based on the interaction of proline with a ninhydrin reagent, forming a pink-red color. The amount of polyphenolic compounds was determined by the Folin-Ciocalteu method. It was found that the ratio of plants grown together significantly affects the accumulation of primary and secondary metabolites in their leaves. Under conditions of mixed planting, more proline and phenolic substances are accumulated in plant leaves compared to monoculture. The maximum proline content, 19.44 ± 0.91 mg/g of dry weight (DW), was observed in the leaves of A. arguta in the experiment combination with a prevailing number of schisandra plants at a ratio of S. chinensis and A. arguta plants of 2:1. In the same combination, the leaves of actinidia contained the highest amount of phenolic compounds (36.87 ± 2.22 mg/g DW). The studied root exudates of the experimental plants had an allelopathic inhibitory effect on the test culture. The exudates caused 12.0 % average decrease in root growth of A. arguta test objects, and 30.0 % average decrease in root growth of S. chinensis test objects compared with the control. This allows us to conclude about the high activity of schisandra’s allelochemicals, which negatively affect the development of actinidia plants. The optimal ratio of plants A. arguta and S. chinensis when grown together is 2 : 1, since a higher concentration of schisandra plants in a container more stress in actinidia plants, expressed as an increased accumulation of phenols and prolines in its leaves. The result of a comparative analysis of the amount of proline and phenolic compounds can be used to assess the mutual influence of plants in mixed plantings to optimize their growing conditions, which confirms the diagnostic significance of these metabolites for indicating the stress state of the studied plants. |
| doi_str_mv | 10.46341/PI2021016 |
| first_indexed | 2025-07-17T12:54:00Z |
| format | Article |
| fulltext |
© The Authors. This content is provided under CC BY 4.0 license.
Plant Introduction, 93/94, 18–26 (2022)
RESEARCH ARTICLE
Diagnostic significance of the synthesis of phenolic compounds and
proline in the leaves of Schisandra chinensis and Actinidia arguta for
the indication of the stress levels of plants under conditions of mixed
plantings
Tetyana Venediktova *, Natalia Zaimenko, Nadiia Skrypchenko
M.M. Gryshko National Botanical Garden, National Academy of Sciences of Ukraine, Tymiryazevska str. 1, 01014 Kyiv, Ukraine;
* tatianaforest3@gmail.com
Received: 29.11.2021 | Accepted: 20.12.2021 | Published online: 21.01.2022
Abstract
The peculiarities of the accumulation of polyphenolic compounds and free proline were investigated in
the leaves of Actinidia arguta and Shisandra chinensis during their cultivation in vegetation containers with
different ratios of the number of plants, namely 1 : 1, 2 : 1, 1 : 2. Monocultural (single-species) planting was
used as a control. The content of free proline in plant leaves was carried out according to the method,
which is based on the interaction of proline with a ninhydrin reagent, forming a pink-red color. The amount
of polyphenolic compounds was determined by the Folin-Ciocalteu method. It was found that the ratio
of plants grown together significantly affects the accumulation of primary and secondary metabolites in
their leaves. Under conditions of mixed planting, more proline and phenolic substances are accumulated
in plant leaves compared to monoculture. The maximum proline content, 19.44 ± 0.91 mg/g of dry weight
(DW), was observed in the leaves of A. arguta in the experiment combination with a prevailing number of
schisandra plants at a ratio of S. chinensis and A. arguta plants of 2:1. In the same combination, the leaves of
actinidia contained the highest amount of phenolic compounds (36.87 ± 2.22 mg/g DW). The studied root
exudates of the experimental plants had an allelopathic inhibitory effect on the test culture. The exudates
caused 12.0 % average decrease in root growth of A. arguta test objects, and 30.0 % average decrease in
root growth of S. chinensis test objects compared with the control. This allows us to conclude about the high
activity of schisandra’s allelochemicals, which negatively affect the development of actinidia plants. The
optimal ratio of plants A. arguta and S. chinensis when grown together is 2 : 1, since a higher concentration
of schisandra plants in a container more stress in actinidia plants, expressed as an increased accumulation
of phenols and prolines in its leaves. The result of a comparative analysis of the amount of proline and
phenolic compounds can be used to assess the mutual influence of plants in mixed plantings to optimize
their growing conditions, which confirms the diagnostic significance of these metabolites for indicating the
stress state of the studied plants.
Keywords: Actinidia arguta, Shisandra chinensis, phenolic compounds, proline, monocultures, mixed plantings, allelopathic activity
https://doi.org/10.46341/PI2021016
UDC [582.678.2 + 582.688.4] : [581.45 : 581.19] : 581.14
Authors’ contributions: T. Venediktova was engaged in preparing and conducting the biochemical analyses, wrote the methodological
part of the research, realized statistical processing of the experimental data, wrote the manuscript, and formulated conclusions.
N. Zaimenko developed the research concept, interpreted the results. N. Skrypchenko wrote the manuscript, interpreted the results,
and formulated conclusions.
Funding: The research has been carried out according to the science program of the Department of Fruit Plants Acclimatization of
the M.M. Gryshko National Botanical Garden of the NAS of Ukraine “Theoretical bases of adaptive introduction for preservation of
biological diversity of fruit plants of world flora”.
Competing Interests: The authors declare no conflict of interest.
https://creativecommons.org/licenses/by/4.0/
https://orcid.org/0000-0002-7419-0703
https://orcid.org/0000-0003-2379-1223
https://orcid.org/0000-0002-1233-9920
Plant Introduction • 93/94 19
Phenolic compounds and proline in the leaves of Schisandra chinensis and Actinidia arguta
Introduction
One of the directions for intensifying
horticulture is the creation of highly productive
plantations with greater technological and
economic efficiency (Egorov, 2013). The main
goal of intensive horticulture is to increase
the productivity and quality of agricultural
products and reduce material costs for
agricultural production. A characteristic
feature of a garden, as an agricultural
phytocenosis, is its instability associated with
the disruption of the trophic relationships of
its components due to the intensive use of
natural resources and technogenic factors
(Popova, 2005). The garden’s biotope is
subjected to a long-term unilateral impact
due to the use of monoculture and typical
agricultural techniques (Kudasov, 1999).
Gardens are monocultural (i.e., single-species)
plantings, with decreased resistance and
productivity of phytocoenoses. Long-term
permanent cultivation of agricultural plants
leads to a progressive deterioration of soil
fertility and soil exhaustion. According to the
Food and Agriculture Organization (FAO), soil
exhaustion is responsible for the loss of 25 %
of the world crop yield (Zhuchenko, 2008).
Numerous studies indicate that the symptoms
of soil exhaustion are more pronounced in
intensive gardens than in organic ones (Manici
et al., 2003).
Diversifying the species composition of
fruit plantations by increasing the biological
diversity of garden phytocoenoses is
possible due to the introduction of rare
fruit plants into the culture, in particular
the hardy kiwi (Actinidia arguta (Siebold et.
Zucc.) Planch. ex Miq.) and Chinese magnolia
vine (Shisandra chinensis (Turcz.) Baill.) –
valuable fruit, medicinal and ornamental
plants. Fruits of A. arguta accumulate a high
content of biologically active substances
such as vitamins C, E, K, polysaccharides,
polyphenols, triterpenoids, and alkaloids,
which have analgesic, antibacterial,
antioxidant, antitumor, hypoglycemic, and
other pharmacological effects (Sun et al.,
2020). Fruits of S. chinensis is a source of
biologically active compounds – vitamins
C, E, and P, saponins, flavonoids, organic
acids, lignans, pectins, and aromatic
substances, that have adaptogenic, tonic,
immunostimulatory, anti-inflammatory,
regenerative, antitumor, and other effects
(Szopa et al., 2016; Nowak et al., 2019). These
plants are widely cultivated in industrial
and farm horticulture of many countries.
Significant introductory studies of A. arguta
and S. chinensis were carried out at the
M.M. Gryshko National Botanical Garden
(NBG) of the National Academy of Sciences
of Ukraine (Skrypchenko & Latocha, 2017).
However, the development of scientific
principles for their introduction into
horticulture and the perfection of cultivation
technology remains crucial.
According to the concept of green
horticulture, the main principle for creating
garden phytocoenoses should be to optimize
their structure by creating multicomponent
mixed plantings, that is, by moving from
monoculture to polyculture (Moroz, 1990).
Interest in “permanent agriculture” reflects
growing attention to mixed plantings that are
inherently more resilient than monocultures.
They can protect and enrich soil ecosystems,
allow plants to form mutually beneficial
combinations, and create a favorable
microclimate (Millner, 2016). With this in
mind, studies of A. arguta and S. chinensis
were realized to assess the possibility of co-
growth of these fruit vines in mixed plantings,
as these plants are characterized by the same
life form and similar agronomic cultivation
requirements.
Since fruit plants not only absorb the
essential mineral elements and organic
compounds but also release various
metabolites into the environment, the success
of the integration of introduced species
into the agrocoenoses depends on their
allelopathic potential and living compatibility
in mixed plantings (Osipova, 1997).
Physiological adaptation of plants to specific
conditions is achieved through physiological
and biochemical mechanisms. Elucidation
of these mechanisms of plant adaptation
to changing environmental conditions is of
great theoretical and practical importance.
Nowadays, it is one of the essential tasks of
ecological plant physiology (Fedulov et al.,
2015). The presence and accumulation of
secondary metabolites reflect the adaptation
strategy of plants created by natural selection
in the course of evolution.
The effect of any abiotic or biotic stress
factor on the plant organism provokes
20 Plant Introduction • 93/94
T. Venediktova, N. Zaimenko, N. Skrypchenko
overproduction of reactive oxygen species
(ROS) and upsets the balance between the
level of ROS and the activity of the antioxidant
defense system (Mittler, 2002; Foyer & Noctor,
2000). An indicator of stress is the amount of
proline, the content of which increases tens
and hundreds of times under conditions of
drought, salinity, high and low temperatures,
and other damaging factors. Proline is the
most abundant compound accumulated by
plants in response to stress (Bassi & Sharma,
1993; Ashraf & Foolad, 2007; Verbruggen
& Hermans, 2008). It is an osmoprotector,
a stabilizer of macromolecules and
membranes, an additional source of energy
and nitrogen, an antioxidant (Kuznetsov &
Shevyakova, 1999).
The dynamics of accumulation of phenolic
compounds, which are important in the
adaptation of the organism to environmental
conditions, can also be used as a criterion
for assessing the adaptive capacity of species
(Zaprometov, 1993; Polyakova & Yershova,
2000; Lattanzio, 2013). Changes in the content
of phenolic compounds depending on the
growing conditions provide the basis of the
ecological stability of natural populations
in the process of evolution and fit into the
general mechanism of plant adaptation to
habitat conditions (Polyakova, 1993). Phenolic
compounds, or polyphenols, are among the
most common secondary metabolites of
vascular plants and are formed in all plants’
cells and tissues (Alscher & Hess, 2017). The
structure of polyphenols is exceptionally
diverse, as are the functions they perform.
They are known to give color to flowers, fruits,
and seeds and participate in plant growth and
development regulation. They are chelators
of heavy metals, regulate the expression
of certain genes, and protect plants from
stress (Bidel et al., 2010; Mierziak et al., 2014).
Many flavonoids are allelochemicals involved
in the formation of rhizobial symbiosis
and mycorrhiza. The mechanism of action
of allelochemical flavonoids is not fully
understood; it is possible that they affect the
auxin signaling of recipient plants and inhibit
the growth of their cells, disrupt the synthesis
of adenosine triphosphoric acid (ATP), induce
the accumulation of ROS and, through calcium
signaling, systemic root death (Mierziak et al.,
2014). The accumulation level of phenolic
compounds, to a certain extent, can serve as a
criterion for the potential resistance of plants
to stress (Zagoskina, 2005).
Laboratory bioanalysis is the first step
to investigate the possible manifestation
of allelopathy in the relationships between
plants, plants and microorganisms, or
plants and insects (Kondratyev, 2017). It
is a necessary tool both for studying the
allelopathic potential of plant or soil extracts
and assessing the activity of extracts in the
purification and identification of allelopathic
compounds. The bioanalysis method supposes
the germination of recipient plants’ seeds
in Petri dishes on filter paper, sand, soil, or
agar. This is a fast method for a large number
of biological repetitions. It can be used to
identify potential allelopathic effects under
controlled laboratory conditions, using the
percentage of germinated seeds of recipient
plants to measure the allelopathic activity of
compounds (Gawronska et al., 2006).
This work aimed to study the accumulation
peculiarities of proline and phenolic
substances in the leaves of A. arguta and
S. chinensis in vegetation experiments and
model their co-growth to optimize their
cultivation.
Material and methods
The research was carried out at the
Department of acclimatization of fruit plants of
the M.M. Gryshko National Botanical Garden,
National Academy of Sciences of Ukraine. The
experiments were carried out under controlled
conditions. The temperature was maintained
within 22 ± 2 ºC, soil moisture – 60 ± 5 %. Plants
of A. arguta ‘Sentyabrskaya’ and S. chinensis
‘Sadovy-1’ were used as experimental objects.
For each experiment combination, 12 two-
year-old vegetatively propagated plants
were planted in the last decade of May in
12-liter containers filled with dark gray forest
light loamy soil in a ratio of 1 : 1, 2 : 1, and 1 : 2;
monocultural plantings of these plants served
as control. The experiments were triplicated.
Samples for biochemical studies were taken
in mid-August. The coefficient of plant
resistance was calculated as the ratio of the
amount of proline or flavonoids in the leaves
of plants of mixed plantings to the amount
of the corresponding substances in plants in
monoculture.
Plant Introduction • 93/94 21
Phenolic compounds and proline in the leaves of Schisandra chinensis and Actinidia arguta
Determination of proline
Determination of the content of free proline
in plant leaves was carried out according to
Bates et al. (1973). This method is based on
the interaction of proline with a ninhydrin
reagent, forming a pink-red color. The content
of free proline was determined in a 2 g
sample of plant material, which was ground
in a mortar with quartz sand and 20 ml of an
aqueous solution of sulfosalicylic acid. After
that, 2 ml of the filtrate were mixed with 2 ml
of acidic ninhydrin and 2 ml of glacial acetic
acid in a test tube with a ground glass stopper.
The mixture was kept for one hour in a boiling
water bath and then cooled. Benzene (4 ml)
was added to the test tubes with the cooled
mixture and vigorously shaken until the orange
color passed into the organic solvent. The
upper colored layer was poured into cuvettes
(20 mm), and the color density of the solution
was measured using a FEK-56M photoelectric
colorimeter. Extinction was determined on a
blue filter with a wavelength of 520 nm. The
amino acid concentration was calculated using
a calibration curve built on standard proline
solutions and expressed in mg/g of dry
weight (DW).
Determination of polyphenolic compounds
The amount of polyphenolic compounds was
determined by the Folin-Ciocalteu method
(Sibgatullina et al., 2011), which is widely used
to analyze herbal preparations. The method is
based on the Folin-Ciocalteu reagent, a mixture
of phosphotungstic and phosphomolybdate
heteropoly acids, which are reduced by
phenolic compounds in an alkaline medium.
During the reaction, polyphenolic compounds
are oxidized by the action of the reagent, and
a blue color appears due to the formation of a
mixture of reduced tungstates and molybdates.
Namely, 0.05 g of raw material samples
were triturated with 1.5 cm3 of 96 % ethanol;
phenolic compounds were extracted for 45
minutes at 45 °C with periodic stirring (every
15 minutes) and subsequent centrifugation for
2 minutes at a rotational speed of 16,000 rpm.
After that, 0.075 cm3 of samples were taken
from the obtained extract, added 0.075 cm3
of the Folin-Ciocalteu reagent, diluted five
times, and stirred. After 3 minutes, 0.15 cm3
of 20 % sodium carbonate solution and 1.2 cm3
of distilled water were added, covered with a
lid, stirred, and left at room temperature. After
one hour, the optical density of the formed
tungsten blue was measured on a Specord-40
spectrophotometer at a wavelength of 725 nm.
The total content of phenolic compounds
was expressed in mg-equivalents of gallic
acid per g of DW, the color intensity of which
was proportional to the amount of phenolic
compounds.
Determination of allelopathic activity
The allelopathic activity of the samples
was determined according to the standard
technique (Grodzinsky, 1991). To obtain root
exudates, donor plants were grown in special
funnels filled with quartz sand (Lobkov &
Konoshina, 2004). After 14 days of vegetation,
samples of aqueous solutions of root exudates
were taken. The resulting aqueous solutions
containing root exudates were evaporated
in a Heidolph Laborota 4002 rotary vacuum
evaporator at 30 °C and residual air pressure
of 30 mbar. The final solution volume was
obtained at the rate of 1 ml from two and ten
donor plants. The concentrated solutions
were stored at below zero temperatures and
used in further studies. An aqueous solution
in a volume of 5 ml was placed in Petri dishes,
into which 50 pieces of watercress (Lepidium
sativum L.) seeds were sown. The dishes were
kept in the dark at room temperature for
three days. After that, the length of L. sativum
seedling roots was measured. The experiments
were carried out in five replicates. Water
served as a control.
Statistical analysis
The significance of differences between
the combinations of the experiment was
established by the dispersion method
according to Fisher’s test and the significance
level of the null hypothesis. The obtained
indicators, determined with a 95 % confidence
interval, are trustworthy due to the high
reliability of the arithmetic mean values (the
calculated Student’s test value significantly
exceeds the table values) and the error
indicator of experience less than 5 %.
Results and discussion
As a result of the studies, it was found that
in the leaves of A. arguta and S. chinensis
grown in a mixed planting, a greater amount
22 Plant Introduction • 93/94
T. Venediktova, N. Zaimenko, N. Skrypchenko
of proline accumulates in comparison with the
monoculture (Table 1).
The maximum amount of proline (21.19 ± 0.8
mg/g DW) was accumulated in S. chinensis
leaves in the experiment combination with
a prevailing ratio of actinidia plants in mixed
planting (with 33 % of S. chinensis and 67 %
of A. arguta). In this combination of mixed
planting, the ratio of proline content in the
leaves of schisandra to the corresponding
value noted for the monoculture (k) was 3.04,
which is the highest observed. The smallest
k value (1.78) was noted in the experiment
combination with 67 % of S. chinensis and 33 %
A. arguta. This indicates that A. arguta plants
have a repressing effect on S. chinensis plants
in mixed plantings.
The amount of proline in A. arguta plants
in the combination S. chinensis (33 %) /
A. arguta (67 %) was 9.53 ± 0.35 mg/g DW.
A similar proline amount was registered
for the control (monoculture) – 9.64 ± 0.41
mg/g DW. In all other combinations of the
vegetation experiments with mixed plantings,
both the amount of proline and the ratio of
the value of proline in mixed plantings were
much higher than the corresponding values
in monoculture. The maximum content of
proline (19.44 ± 0.91 mg/g DW) was observed in
A. arguta in the experiment combination with
the prevailing numbers of S. chinensis plants,
with 67 % of S. chinensis and 33 % A. arguta,
and with k = 2.02. The smallest coefficient
(k = 0.99) was obtained in the combination
S. chinensis (33 %) / A. arguta (67 %). Thus,
with an increase in the number of S. chinensis
plants in a vegetation container, A. arguta
plants experience more severe stress (Table 1).
A similar relationship was noted for S. chinensis
with an increase of A. arguta plants.
Many plant species naturally accumulate
proline as the main organic osmolytes
when subjected to various abiotic stresses.
Therefore, this compound, which is effectively
involved in the mechanisms of ensuring
plant resistance to stress, particularly plays
an adaptive role in osmotic adaptation and
protects subcellular structures in plants under
stress conditions (Verbruggen & Hermans,
2008; Chutia & Borah, 2012). The response
of fennel plants (Foeniculum vulgare Mill.)
to environmental constraints was a marked
increase in proline content in its leaves (Zali &
Ehsanzadeh, 2018).
The study of the features of the
accumulation of phenols in the leaves of
A. arguta and S. chinensis showed that their
amount in S. chinensis leaves increases in
mixed plantings with the prevailing number of
A. arguta plants (Fig. 1).
The maximum accumulation of phenolic
compounds (36.87 ± 2.22 mg/g DW) in
S. chinensis leaves was observed in the
combination S. chinensis (67 %) / A. arguta
(33 %), where the ratio of phenols in the leaves
of plants in mixed plantings to the amount of
phenols in monoculture plants was the highest
(k = 2.09). The ratio was much lower (k = 1.12)
in the combinations of S. chinensis (50 %) /
A. arguta (50 %) and S. chinensis (33 %) /
A. arguta (67 %). In leaves of A. arguta in
monoculture, and in combinations S. chinensis
(33 %) / A. arguta (67 %), the phenol content
differed insignificantly (24.88 ± 1.43 mg/g DW
and 23.82 ± 1.47 mg/g DW, respectively). In
the experiment combinations S. chinensis
Nr Experiment combination
S. chinensis A. arguta
mg/g DW k mg/g DW k
A S. chinensis (control) 6.98 ± 0.36
A A. arguta (control) 9.64 ± 0.41
B S. chinensis (50 %) / A. arguta (50 %) 19 ± 0.95 2.72 13.57 ± 1.9 1.41
C S. chinensis (33 %) / A. arguta (67 %) 21.19 ± 0.81 3.04 9.53 ± 0.35 0.99
D S. chinensis (67 %) / A. arguta (33 %) 12.4 ± 0.64 1.78 19.44 ± 0.91 2.02
LSD 0.95 1.88 1.65
Table 1. Accumulation of proline in Actinidia arguta and Shisandra chinensis plants and their stress resistance
coefficient in monocultures and mixed plantings.
Note. DW – dry weight; k – stress resistance coefficient; LSD – least significant difference.
Plant Introduction • 93/94 23
Phenolic compounds and proline in the leaves of Schisandra chinensis and Actinidia arguta
(50 %) / A. arguta (50 %) and S. chinensis
(33 %) / A. arguta (67 %), an accumulation of
a higher content of phenolic compounds in
A. arguta leaves was observed. The highest
coefficient of resistance to stress, k = 1.22, was
recorded in the experiment combination with
the prevailing number of S. chinensis plants,
namely S. chinensis (67 %) / A. arguta (33 %).
Thus, it was found that the content of
secondary metabolites of phenolic nature
in the leaves of plants changes depending
on the ratio of experimental plants in the
compositions of mixed plantings (Fig. 1).
The research results revealed interspecific
competition of plants in their co-cultivation,
which leads to stress and can be manifested
in relation to mineral nutrients, soil moisture,
or be caused by root secretions of plants. The
concentration and amount of biosynthesized
phenolic compounds increase responding to
stress factors. A similar reaction of plants is
reported in several publications. In particular,
the water stress increased the amount of
the phenolic compounds in the vegetative
organs of Hypericum brasiliense Choisy
(Abreu & Mazzafera, 2005). Similarly, water-
deficit stress increased the productivity of
chlorogenic acid, catechin and epicatechin in
hawthorn species (Kirakosyan et al., 2004).
To compare the allelopathic potential
of the experimental plants, we studied the
allelopathic activity of their root exudates by
the biotesting method using the L. sativum as
a test object (Fig. 2).
The allelopathic activity of the root
exudates of A. arguta and S. chinensis plants
was established. Their noticeable phytotoxic
effect on the growth of roots of L. sativum
seedlings was found. Compared with the
control, the exudate of A. arguta suppressed
L. sativum roots growth by 12.0 % and
S. chinensis – by 30.0 %. A decrease in the
growth of L. sativum roots under the influence
of aqueous solutions of root exudates indicates
the presence of dissolved organic substances
in the soil, which inhibit the development
of test objects. As noted above, S. chinensis
exhibits a stronger inhibitory effect on the
test culture compared to A. arguta. These
results confirm the previously obtained
data that intravital secretions (leaf and
root exudates) of S. chinensis and A. arguta
contain allelopathically active compounds –
allelochemicals, which have a depressing effect
on seed germination and growth of seedlings
of various test objects (Osipova, 1997).
Conclusions
The results of the vegetation experiment
indicate a pronounced interaction between
A. arguta and S. chinensis crops during
joint growth. The percentage of plants in
model experiments significantly affects the
accumulation of primary and secondary
metabolites in leaves.
Figure 1. Accumulation of phenolic substances in
plants of Actinidia arguta and Shisandra chinensis in
monocultures and mixed plantings: A – A. arguta
(control); B – S. chinensis (50 %) / A. аrguta (50 %);
C – S. chinensis (33 %) / A. arguta (67 %); D – S. chinensis
(67 %) / A. arguta (33 %).
Figure 2. Allelopathic activity of root secretions of
Actinidia arguta and Shisandra chinensis (bioassay
– root growth of Lepidum sativum), percentage to
control.
24 Plant Introduction • 93/94
T. Venediktova, N. Zaimenko, N. Skrypchenko
The result of a comparative analysis of the
amount of proline and phenolic compounds
can be used to assess the mutual influence
of plants in mixed plantings to optimize their
growing conditions, which confirms the
diagnostic significance of these metabolites
for indicating the stress state of the studied
plants.
The inhibitory effect of S. chinensis and
A. arguta root secretions on L. sativum
indicates their allelopathic activity, more
pronounced in S. chinensis.
Based on our results, it is concluded that
monoculture cultivation is the most optimal
for both studied species. However, for the co-
growth of A. arguta and S. chinensis, planting
in the ratio of 2 : 1 is the best.
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Діагностичне значення синтезу фенольних сполук і проліну в листках Schisandra
chinensis та Actinidia arguta для індикації рівня стресу рослин в умовах змішаних
насаджень
Тетяна Венедиктова *, Наталія Заіменко, Надія Скрипченко
Національний ботанічний сад імені М.М. Гришка НАН України, вул. Тимірязєвська, 1, Київ, 01014,
Україна; * tatianaforest3@gmail.com
Досліджено особливості накопичення поліфенольних сполук та вільного проліну в листках Actinidia
arguta та Shisandra chinensis при їх вирощуванні в вегетаційних дослідах з різним співвідношенням
кількості рослин в контейнерах, а саме 1 : 1, 2 : 1, 1 : 2. В якості контролю використовували одновидові
посадки рослин. Вміст вільного проліну в листках рослин здійснювали за методикою, яка заснована
на взаємодії проліну з реактивом нінгідрину, утворюючи рожево-червоне забарвлення. Кількість
поліфенольних сполук визначали методом Фоліна-Чіокальтеу. Встановлено, що співвідношення
рослин в контейнерах істотно впливає на накопичення первинних і вторинних метаболітів у їх
https://doi.org/10.1080/13549839.2016.1272560
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https://doi.org/10.1007/s11101-016-9470-4
https://doi.org/10.1007/s11101-016-9470-4
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https://doi.org/10.1016/j.indcrop.2017.10.020
26 Plant Introduction • 93/94
T. Venediktova, N. Zaimenko, N. Skrypchenko
листках. За умов змішаної посадки в листках рослин накопичувалося більше проліну та фенольних
речовин порівняно з монокультурою. Максимальний вміст проліну 19,44 ± 0,91 мг/г сухої маси
(СМ) спостерігався в листках A. arguta у варіанті з переважаючою кількістю рослин лимонника при
співвідношенні рослин S. chinensis і A. arguta 2 : 1. У цьому ж варіанті найбільша кількість фенольних
сполук (36,87 ± 2,22 мг/г СМ) була у листках актинідії. Досліджувані кореневі ексудати S. chinensis і
A. arguta мали інгібуючу алелопатичну дію на тест-культуру. Ексудати A. arguta викликали середнє
зниження росту коренів тест-об’єктів на 12,0 %, а S. chinensis – на 30,0 % порівняно з контролем.
Це дозволяє зробити висновок про високу активність алелохімікатів лимонника, які негативно
впливають на ріст рослин актинідії. Оптимальне співвідношення рослин A. arguta і S. chinensis при
спільному вирощуванні становить 2 : 1, оскільки вища концентрація рослин лимонника в контейнері
посилює стрес у рослин актинідії, що виражається в підвищеному накопиченні фенолів і проліну
в її листках. Результат порівняльного аналізу кількості проліну та фенольних сполук може бути
використаний для оцінки взаємного впливу рослин у змішаних насадженнях з метою оптимізації
умов їх вирощування, що підтверджує діагностичну значущість цих метаболітів для індикації
стресового стану рослин.
Ключові слова: Actinidia arguta, Shisandra chinensis, фенольні сполуки, пролін, монокультура, змішані насадження, алелопатична
активність
|
| id | oai:ojs2.plantintroduction.org:article-1597 |
| institution | Plant Introduction |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2025-07-17T12:54:00Z |
| publishDate | 2022 |
| publisher | M.M. Gryshko National Botanical Garden of the NAS of Ukraine |
| record_format | ojs |
| resource_txt_mv | wwwplantintroductionorg/bd/044f27ac76db92cc95addcf7feaa04bd.pdf |
| spelling | oai:ojs2.plantintroduction.org:article-15972023-08-26T20:38:56Z Diagnostic significance of the synthesis of phenolic compounds and proline in the leaves of Schisandra chinensis and Actinidia arguta for the indication of the stress levels of plants under conditions of mixed plantings Діагностичне значення синтезу фенольних сполук і проліну в листках Schisandra chinensis та Actinidia arguta для індикації рівня стресу рослин в умовах змішаних насаджень Venediktova, Tetyana Zaimenko, Natalia Skrypchenko, Nadiia The peculiarities of the accumulation of polyphenolic compounds and free proline were investigated in the leaves of Actinidia arguta and Shisandra chinensis during their cultivation in vegetation containers with different ratios of the number of plants, namely 1 : 1, 2 : 1, 1 : 2. Monocultural (single-species) planting was used as a control. The content of free proline in plant leaves was carried out according to the method, which is based on the interaction of proline with a ninhydrin reagent, forming a pink-red color. The amount of polyphenolic compounds was determined by the Folin-Ciocalteu method. It was found that the ratio of plants grown together significantly affects the accumulation of primary and secondary metabolites in their leaves. Under conditions of mixed planting, more proline and phenolic substances are accumulated in plant leaves compared to monoculture. The maximum proline content, 19.44 ± 0.91 mg/g of dry weight (DW), was observed in the leaves of A. arguta in the experiment combination with a prevailing number of schisandra plants at a ratio of S. chinensis and A. arguta plants of 2:1. In the same combination, the leaves of actinidia contained the highest amount of phenolic compounds (36.87 ± 2.22 mg/g DW). The studied root exudates of the experimental plants had an allelopathic inhibitory effect on the test culture. The exudates caused 12.0 % average decrease in root growth of A. arguta test objects, and 30.0 % average decrease in root growth of S. chinensis test objects compared with the control. This allows us to conclude about the high activity of schisandra’s allelochemicals, which negatively affect the development of actinidia plants. The optimal ratio of plants A. arguta and S. chinensis when grown together is 2 : 1, since a higher concentration of schisandra plants in a container more stress in actinidia plants, expressed as an increased accumulation of phenols and prolines in its leaves. The result of a comparative analysis of the amount of proline and phenolic compounds can be used to assess the mutual influence of plants in mixed plantings to optimize their growing conditions, which confirms the diagnostic significance of these metabolites for indicating the stress state of the studied plants. Досліджено особливості накопичення поліфенольних сполук та вільного проліну в листках Actinidia arguta та Shisandra chinensis при їх вирощуванні в вегетаційних дослідах з різним співвідношенням кількості рослин в контейнерах, а саме 1 : 1, 2 : 1, 1 : 2. В якості контролю використовували одновидові посадки рослин. Вміст вільного проліну в листках рослин здійснювали за методикою, яка заснована на взаємодії проліну з реактивом нінгідрину, утворюючи рожево-червоне забарвлення. Кількість поліфенольних сполук визначали методом Фоліна-Чіокальтеу. Встановлено, що співвідношення рослин в контейнерах істотно впливає на накопичення первинних і вторинних метаболітів у їх листках. За умов змішаної посадки в листках рослин накопичувалося більше проліну та фенольних речовин порівняно з монокультурою. Максимальний вміст проліну 19,44 ± 0,91 мг/г сухої маси (СМ) спостерігався в листках A. arguta у варіанті з переважаючою кількістю рослин лимонника при співвідношенні рослин S. chinensis і A. arguta 2 : 1. У цьому ж варіанті найбільша кількість фенольних сполук (36,87 ± 2,22 мг/г СМ) була у листках актинідії. Досліджувані кореневі ексудати S. chinensis і A. arguta мали інгібуючу алелопатичну дію на тест-культуру. Ексудати A. arguta викликали середнє зниження росту коренів тест-об’єктів на 12,0 %, а S. chinensis – на 30,0 % порівняно з контролем. Це дозволяє зробити висновок про високу активність алелохімікатів лимонника, які негативно впливають на ріст рослин актинідії. Оптимальне співвідношення рослин A. arguta і S. chinensis при спільному вирощуванні становить 2 : 1, оскільки вища концентрація рослин лимонника в контейнері посилює стрес у рослин актинідії, що виражається в підвищеному накопиченні фенолів і проліну в її листках. Результат порівняльного аналізу кількості проліну та фенольних сполук може бути використаний для оцінки взаємного впливу рослин у змішаних насадженнях з метою оптимізації умов їх вирощування, що підтверджує діагностичну значущість цих метаболітів для індикації стресового стану рослин. M.M. Gryshko National Botanical Garden of the NAS of Ukraine 2022-01-20 Article Article application/pdf https://www.plantintroduction.org/index.php/pi/article/view/1597 10.46341/PI2021016 Plant Introduction; No 93/94 (2022); 18-26 Інтродукція Рослин; № 93/94 (2022); 18-26 2663-290X 1605-6574 10.46341/PI93-94 en https://www.plantintroduction.org/index.php/pi/article/view/1597/1525 Copyright (c) 2022 Tetyana Venediktova, Natalia Zaimenko, Nadiia Skrypchenko http://creativecommons.org/licenses/by/4.0 |
| spellingShingle | Venediktova, Tetyana Zaimenko, Natalia Skrypchenko, Nadiia Діагностичне значення синтезу фенольних сполук і проліну в листках Schisandra chinensis та Actinidia arguta для індикації рівня стресу рослин в умовах змішаних насаджень |
| title | Діагностичне значення синтезу фенольних сполук і проліну в листках Schisandra chinensis та Actinidia arguta для індикації рівня стресу рослин в умовах змішаних насаджень |
| title_alt | Diagnostic significance of the synthesis of phenolic compounds and proline in the leaves of Schisandra chinensis and Actinidia arguta for the indication of the stress levels of plants under conditions of mixed plantings |
| title_full | Діагностичне значення синтезу фенольних сполук і проліну в листках Schisandra chinensis та Actinidia arguta для індикації рівня стресу рослин в умовах змішаних насаджень |
| title_fullStr | Діагностичне значення синтезу фенольних сполук і проліну в листках Schisandra chinensis та Actinidia arguta для індикації рівня стресу рослин в умовах змішаних насаджень |
| title_full_unstemmed | Діагностичне значення синтезу фенольних сполук і проліну в листках Schisandra chinensis та Actinidia arguta для індикації рівня стресу рослин в умовах змішаних насаджень |
| title_short | Діагностичне значення синтезу фенольних сполук і проліну в листках Schisandra chinensis та Actinidia arguta для індикації рівня стресу рослин в умовах змішаних насаджень |
| title_sort | діагностичне значення синтезу фенольних сполук і проліну в листках schisandra chinensis та actinidia arguta для індикації рівня стресу рослин в умовах змішаних насаджень |
| url | https://www.plantintroduction.org/index.php/pi/article/view/1597 |
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