Прояв фенотипової пластичності листків псамофіту Corynephorus canescens при затопленні
The results of the study of the leaf structure in psammophyte Corynephorus canescens, which grew under controlled conditions and flooding using the methods of light microscopy, scanning electron microscopy, and laser confocal microscopy, are presented. This study revealed common and distinctive sign...
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2021
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| description | The results of the study of the leaf structure in psammophyte Corynephorus canescens, which grew under controlled conditions and flooding using the methods of light microscopy, scanning electron microscopy, and laser confocal microscopy, are presented. This study revealed common and distinctive signs of morphological and anatomical parameters of C. canescens leaves in the phase of vegetative growth. Among the common features were the shape and size of the leaf laminas, hypostomatic type of the leaf, isolateral structure of the parenchyma, the thick-walled epidermis, and the bilayered hypodermis. Among the distinctive features were the signs of the destruction of cells in the photosynthetic parenchyma, change in their shape with the formation of protuberances at the cells’ poles, and almost doubling area of the aerenchyma in C. canescens leaves under flooding conditions. Scanning electron microscopy showed the similarity of ultrastructure and density of trichomes on the adaxial surface, excepting the formation of cuticular wax structures on the epidermal surface of the leaves in flooded plants. The subcellular localization of silicon inclusions was studied for the first time. The presence of amorphous and small crystalline silicon inclusions in the periclinal walls of the main epidermal cells and amorphous silicon inclusions in leaf trichomes was established. An increase in the relative silicon content along the trichomes in the leaves’ epidermis after flooding was revealed. It was assumed that the phenotypic plasticity of C. canescens, is realized through the increasing area of aerenchyma in leaves and increasing silicon content in trichomes. Such plasticity helps to optimize both the oxygen balance of plants and water balance in flooded plants, thus increasing the species’ resistance to prolonged flooding. |
| doi_str_mv | 10.46341/PI2021011 |
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© The Authors. This content is provided under CC BY 4.0 license.
Plant Introduction, 91/92, 24–35 (2021)
RESEARCH ARTICLE
Appearance of phenotypic plasticity of leaves in psammophyte
Corynephorus canescens during flooding
Оlena Nedukha
Department of Cell Biology and Anatomy, M.G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine,
Tereschenkivska str. 2, 01601 Kyiv, Ukraine; o.nedukha@hotmail.com
Received: 13.10.2021 | Accepted: 15.11.2021 | Published online: 21.11.2021
Abstract
The results of the study of the leaf structure in psammophyte Corynephorus canescens, which grew under
controlled conditions and flooding using the methods of light microscopy, scanning electron microscopy,
and laser confocal microscopy, are presented. This study revealed common and distinctive signs of
morphological and anatomical parameters of C. canescens leaves in the phase of vegetative growth. Among
the common features were the shape and size of the leaf laminas, hypostomatic type of the leaf, isolateral
structure of the parenchyma, the thick-walled epidermis, and the bilayered hypodermis. Among the
distinctive features were the signs of the destruction of cells in the photosynthetic parenchyma, change
in their shape with the formation of protuberances at the cells’ poles, and almost doubling area of the
aerenchyma in C. canescens leaves under flooding conditions. Scanning electron microscopy showed
the similarity of ultrastructure and density of trichomes on the adaxial surface, excepting the formation
of cuticular wax structures on the epidermal surface of the leaves in flooded plants. The subcellular
localization of silicon inclusions was studied for the first time. The presence of amorphous and small
crystalline silicon inclusions in the periclinal walls of the main epidermal cells and amorphous silicon
inclusions in leaf trichomes was established. An increase in the relative silicon content along the trichomes
in the leaves’ epidermis after flooding was revealed. It was assumed that the phenotypic plasticity of
C. canescens, is realized through the increasing area of aerenchyma in leaves and increasing silicon content
in trichomes. Such plasticity helps to optimize both the oxygen balance of plants and water balance in
flooded plants, thus increasing the species’ resistance to prolonged flooding.
Keywords: Corynephorus canescens, leaf epidermis, flooding, aerenchyma
https://doi.org/10.46341/PI2021011
UDC 581.45 : 58.032 : 549.514.5
Funding: This research was supported partly by the theme N467 “Cellular and molecular mechanisms of phenotypic plasticity of
psammophytes and heliophytes in contrast conditions of water regime” of the National Academy of Sciences of Ukraine.
Competing Interests: The author claims that there is no conflict of interest regarding the research, authorship and/or publication
of this article.
Introduction
The plants growing on sandy soils in steppe
and desert areas and on the sandy shores of
seas, rivers, and lakes are psammophytes.
The study of morphological and anatomical
features of psammophytes is important for
the understanding of the plant adaptation
and response to various external influences
(Hameed et al., 2009; Elhalim et al., 2016). It
is known that the vegetation of coastal sandy
dunes is constantly exposed to repeated
https://creativecommons.org/licenses/by/4.0/
https://orcid.org/0000-0001-8405-453X
Plant Introduction • 91/92 25
Phenotypic plasticity of Corynephorus canescens leaves during flooding
environmental stresses, which directly affects
the survival and distribution of plant species.
The most significant ecological stresses for
psammophytes are drought, salinity, solar
radiation, high intensity of light, low nutrient
availability, and soil instability (Grigore &
Toma, 2007; Bagousse-Pinguet et al., 2013).
Mentioned factors closely interact as abiotic
modulators of leaf adaptation and are
directly regulated by primary stress signals,
which trigger the appropriate structural and
functional adaptation mechanism required
for plant survival (Catoni & Gratani, 2013).
Psammophytes growing on sandy dunes have
developed mechanisms of resistance and
adaptation to environmental changes (Ashraf
& Harris, 2013; Ruocco et al., 2014; Futorna
et al., 2017).
The morphological and anatomical
features of the psammophytes’ adaptation to
environmental conditions are: the decrease of
the leaves’ size, the formation of water-storing
parenchyma, changes in the foliar vascular
system size, leaf laminas twisting, changes
in stomatal density, increased indumentum,
thickened cuticles, and synthesis of calcium
inclusions in cells and intercellular spaces
(Hameed et al., 2009; Elhalim et al., 2016;
Jianu et al., 2021). Recently, silicon inclusions
were found to be associated with plant
resistance to changes in cell water balance.
Notably, during drought, silicon enhances the
suberinization of the sclerenchyma of rice
roots and lignification of the foliar epidermis of
heliophyte Phragmites australis (Cav.) Trin. ex
Steud. (Fleck et al., 2011; Nedukha & Kordyum,
2019). Psammophytes, in addition to the above-
mentioned environmental stresses, may be
affected by short-term or prolonged flooding.
However, the adaptability of psammophytes to
flooding has not been investigated.
Water is one of the main factors supporting
the growth, development, and productivity
of such psammophyte as Corynephorus
canescens (L.) P. Beauv. (Poaceae). Considering
that aerenchyma and epidermis of the leaves
are principal tissues regulating the water
balance of the plant (Kerstiens, 2006; Hansen
et al., 2007), our work aimed to investigate
in C. canescens: 1) the general anatomical
structure of the leaves; 2) the epidermal
ultrastructure of the leaves under different
water supply conditions; 3) the location and
content of silicon in the foliar epidermis to
understand the role of this element in the
plant adaptation to flooding.
Material and methods
The leaves of two study variants of
C. canescens (grown on slightly moist sandy
soil and flooded for 21 days) were investigated.
At the end of May, the turf of 30-days plants
of C. canescens, which grew on sandy soil
(Fig. 1 A), was planted in cups filled with river
sand. In total, six cups containing 17 plants per
cup were obtained. Three cups were flooded
(Fig. 1 B) to the level of plants’ hypocotyl. The
other three cups with plants were not flooded
and served a control. The duration of daylight
for all six sampling series was from 16 h 8 m
to 16 h 26 m, with daytime temperature 22–
28 °C and nighttime temperature 20–23 °C.
The control plants were watered every two
days with 15 ml of settled running water.
Standard biochemical methods were applied
to determine and control the relative water
content of the soil (Ermakov, 1982).
The histological slides were prepared
following standard methods for cytological
studies of the leaf structure under the
light and scanning electron microscopy
(Pausheva, 1988). For the light microscopy,
the samples were fixed with a mixture of 2 %
paraformaldehyde and 2 % glutaraldehyde
(1 : 1, vol.) for 12 h, dehydrated with alcohol
and acetone, and saturated with a mixture
of epoxy resins (epon-araldite) (Pausheva,
1988). Sections (10 μm thick) were stained
with 0.05 % toluidine blue O and 0.01 %
safranin following Schmid (1980), and then
studied with a light microscope Carl Zeiss NF
(Germany). For scanning electron microscopy,
samples were fixed in a mixture of aldehydes
and dehydrated, fixed on special stubs,
sputter-coated with gold, and studied under
an electron microscope Jeol JSM 6060 LA
(Japan). For cytochemical studies of the
localization of silicon inclusions, clippings
from the middle part of the leaf laminas were
examined on a laser confocal microscope
Zeiss LSM5 (Germany) using an excitation
wavelength of 480 nm and emission of 530 nm.
The fluorescence intensity of silicon was
determined by the Pascal program (Zeiss,
Germany).
26 Plant Introduction • 91/92
O. Nedukha
BA
Figure 1. Study variants of Corynephorus canescens: A – general view of plants grown 30 days in semi-sandy
soil, the phase of vegetative growth; B – plants transplanted into the sandy soil and serving a control
variant (the top row) and flooded plants serving an experiment variant (the bottom row).
Results
The leaves of C. canescens in both sampling
series were characterized by identical
morphological structure (Fig. 1). In particular,
leaves were sessile, simple, linear, twisted,
hard, gray-green, with a rough surface and
parallel venation. Their size in the rolling
state was 130.0 ± 3.1 mm on the long axis and
2.7 ± 0.3 mm on the short axis. Their size on
the short axis in the unwind state of leaf
blade was from 7.3 ± 1 mm to 12.0 ± 2 mm. The
anatomical structure of the leaves showed
some differences between the variants that
are described below.
Light microscopy
Control variant. The plants of C. canescens
grown without flooding (control conditions)
had twisted leaves with hippocrepiform
cross-sections (Fig. 2 A). The upper (adaxial)
leaf surface was uniform. The lower (abaxial)
leaf surface had hollows and protrusions
(Fig. 2 A, B). The protrusions were located near
the vascular bundles. The sizes of epidermis
and mesophyll cells differed among the
surfaces (Table 1).
The thickness of the leaf lamina in the
area of protrusions ranged from 237 ± 13 μm
(in the center of the lamina) to 200 ± 11 μm (in
the periphery of leaf lamina). The thickness
of the leaf lamina in the depressions was
from 150 ± 10 μm to 200 ± 13 μm, respectively.
Leaf laminas were hypostomatic with the
isolateral structure of the parenchyma, thick-
walled epidermis, and bilayered hypodermis
containing inclusions.
The cells of the adaxial epidermis were
thick-walled. The thickness of the cell walls
in the adaxial epidermis was 3.2 ± 0.2 μm, the
height of the cells was 13.5 ± 1.1 μm, and the
width was 16.0 ± 1.3 μm.
Behind the epidermis, depending on the cut
and the surface area, were present one or two
layers of thick-walled hypodermal cells. The
thickness of the cell walls in the hypodermal
cells was 2.7 ± 0.1 μm; the average cell diameter
was 7.1 ± 0.1 μm. Occasionally, in hypodermal
cells, the inclusions of various forms were
visible.
Elongated, compacted, thin-walled
mesophyll cells were located behind the
hypodermis. The height of such cells was
27.3 ± 3.1 μm, width – 10.7 ± 0.7 μm. The average
number of chloroplasts per section of the
mesophyll cell was 11 ± 1.2 units. Four-six layers
of rounded (diameter 12.2 ± 1.1 μm) and slightly
elongated (long axis – 15.4 ± 2.1 μm, short axis
– 11.2 ± 1.3 μm) cells were situated below the
initial mesophyll.
Between the mesophyll layers, small air
cavities representing aerenchyma were visible.
It extended to the abaxial epidermis. The
Plant Introduction • 91/92 27
Phenotypic plasticity of Corynephorus canescens leaves during flooding
aerenchyma area was 21.1 ± 0.7 % of the total
area of the leaf lamina cross-section.
The cells of the abaxial epidermis were
almost rectangular. The average height of the
cells was 11.7 ± 1.1 μm, the width – 10.8 ± 0.4 μm.
The cell walls in abaxial epidermis were thinner
than in the adaxial one, and their thickness
was about 0.9 ± 0.1 μm only. Between the cells
of the abaxial epidermis, the guard cells of the
stomata were present. On the surface of the
abaxial epidermis, long transparent trichomes
were visible.
The cross-section of the leaf lamina had
seven vascular bundles: one large in the center
(diameter was about 52 μm) and six smaller
(diameter was about 10 μm) on the sides.
Flooded variant. 21-day flooding of
C. canescens planted on sandy soil (with
soil moisture of 70.6 ± 2.3 %) did not affect
the morphology of leaf laminas but caused
changes in their anatomical structure. After
14 days of flooding, every fifth leaf changed its
color from green to brown. After 21 days, the
number of brown dead leaves increased.
We used for the anatomical investigations
the leaves that remained green after
21-day flooding. We did not reveal the
changes in the structure of epidermal and
subepidermal (hypodermal) cells. However,
we noted the changes in the size and shape
of mesophyll cells (Table 1; Fig. 2 B, C). Cells
of the upper mesophyll layer changed their
shape and density. The shape of mesophyll
cells changed from elongated to uneven
with the protuberances on cell poles. The
density of mesophyll cells decreased. The
large intercellular spaces (aerenchyma)
emerged between the cells of the upper
mesophyll layer and extended from adaxial
to abaxial epidermis. Hence, the area of
the aerenchyma increased compared to the
A B
C D
Figure 2. Cross-sections of the middle part of Corynephorus canescens leaf lamina: A, B – control variant;
C, D – after 21 days of flooding. abe – abaxial epidermis; ade – adaxial epidermis; aer – aerenchyma;
m – mesophyll.
28 Plant Introduction • 91/92
O. Nedukha
control, and occupied 35.0 ± 2.9 % of the
cross-sectional area of the leaf lamina.
Scanning electron microscopy
Control variant. The plants of C. canescens,
which grew in the control conditions, had
irregular epidermis with protrusions of the
periclinal walls of the main epidermal cells
located along adaxial surface of the leaf
lamina. Whole epidermal cells were covered
with a solid layer of cuticular wax depositions.
The apexes of trichomes were oriented in
one direction – from the leaf base to its tip
(Fig. 3 A). Trichomes were short, with the
rectangular or slightly oval shape of the base
Parameter Control (soil moisture –
42.7 ± 2.1 %)
21-day flooding (soil
moisture – 70.6 ± 2.3 % *)
Leaf shape Linear Linear
Leaf sizes:
long axis, mm 13.0 ± 1.3 13.0 ± 1.1
short axis (in rolling state), mm 2.7 ± 0.3 2.8 ± 0.2
Area of aerenchyma on the leaf cross-section, % 21.1± 0.7 35.0 ± 2.9 *
Sizes of adaxial epidermis cells:
width, µm 16.0 ± 1.3 16.0 ± 1.3
height, µm 13.5 ± 1.1 13.7 ± 1.1
Diameter of hypodermal cells, µm 7.1 ± 0.1 7.1 ± 0.2
Sizes of mesophyll cells:
first layer
width, µm 10.7 ± 0.7 12.5 ± 1.1
height, µm 27.3 ± 3.1 17.0 ± 1.3 *
second–sixth layers
width, µm 11.2 ± 1.1 10.2 ± 1.4
height, µm 15.4 ± 2.1 13.4 ± 2.1
Sizes of abaxial epidermis cells:
width, µm 10.8 ± 0.4 10.8 ± 0.4
height, µm 11.7 ± 1.1 11.7 ± 1.1
Density of trichomes in adaxial epidermis per 1 mm2 (SEM) 314 ± 27 342 ± 31
Size of trichomes in adaxial epidermis:
basal width, µm 15.3 ± 2.2 15.7 ± 1.7
height, µm 60.0 ± 1.3 57.1 ± 3.1
Density of trichomes in abaxial epidermis per 1 mm2 (SEM) 900 ± 27 879 ± 71
Size of trichomes in abaxial epidermis:
basal width, µm 13.0 ± 1.2 13.0 ± 1.2
height, µm 76.0 ± 7.1 73.0 ± 7.9
Content of silicon inclusions in the cells of adaxial
epidermis (LCM):
at the base of trichomes, in relative units 98.0 ± 3.1 89.7 ± 3.5
along the trichome, in relative units 64.7 ± 0.3 105.7 ± 7.3 *
in the main epidermal cells, in relative units 51.5 ± 0.9 46.3 ± 1.2
Table 1. Physiological and structural parameters of Corynephorus canenscens leaves.
Note. * – р < 0.05 (two-way ANOVA); SЕМ – scanning electron microscopy data; LCM – laser confocal
microscopy data.
Plant Introduction • 91/92 29
Phenotypic plasticity of Corynephorus canescens leaves during flooding
and a hook-shaped apex (Fig. 3 B). The length
of trichomes was 60.0 ± 1.3 μm, the base width
– 15.3 ± 2.2 μm, the trichomes’ middle part
width – 30.0 ± 2.2 μm, the length of trichomes’
hook – 10.0 ± 1.3 μm. The trichomes’ surface
was slightly smooth, similar to the surface of
the surrounding cells. The average density of
trichomes on the upper epidermis of the leaf
was 314 ± 27 trichomes per 1 mm2 (Table 1).
Trichomes were situated in parallel rows; the
distance (in one row) between them varied
from 50 to 100 μm.
The abaxial surface of the leaf was
characterized by zonality (Fig. 4 A, B). It had
alternated zones with longitudinally oriented
trichomes and stomata. The width of each
zone was about 100 μm. Trichome areas
raised above the stomata zones and covered
them. The length of saber-like trichomes
was 76.0 ± 7.1 μm, the width of the trichomes’
base was 13.0 ± 1.2 μm. The average density of
trichomes was 900 ± 27 trichomes per 1 mm2.
Trichomes were placed in parallel rows, and
the distance (in one row) between trichomes
varied from 50 to 100 μm. The stomata
raised above the epidermal surface and were
arranged in rows. The average size of the
stomatal guard cell was 30 ± 1.5 × 6.1 ± 1.1 μm.
The distance between the stomata in one
row varied from 80 to 145 μm; the distance
between stomatal rows was up to 20 μm. The
type of stomata cannot be clearly determined
due to the presence of continuous cuticular-
waxy inclusions on the cells surrounding the
stomata.
Flooded variant. Corynephorus canescens
plants, which were flooded for 21 days, showed
ultrastructure of the epidermis identical with
control samples (Fig. 3 C, D). In particular,
short trichomes were arranged in the same
longitudinal rows. The shape and size of
trichomes were similar to those in the control
samples (Table 1).
Figure 3. Ultrastructure of the adaxial surface of the leaves in Corynephorus canescens grown under control
(A, B) and flooded (C, D) conditions.
A B
C D
30 Plant Introduction • 91/92
O. Nedukha
A B
C D
Figure 4. Ultrastructure of the abaxial surface of the leaves in Corynephorus canescens grown under control
(A, B) and flooded (C, D) conditions.
On the adaxial leaf surface, only certain
features were found in flooded plants. In
particular, we detected verrucous structures
on the surface of the main epidermal cells
and at the base of trichomes (Fig. 3 D). The
average density of trichomes on the adaxial
leaf surface was 342 ± 31 per 1 mm2. Trichomes
were also arranged in rows; the distance (in
one row) between them varied from 50 to
120 μm. Trichomes were short, similar in shape
to those in the control variant, slightly oval
at the base (top view), and had hook-shaped
tips (well-visible in the side view). Trichomes’
length was 57.0 ± 3.1 μm, the width of the
trichomes’ base was 15.0 ± 1.7 μm; the width
of trichomes’ middle part was 30 ± 2.2 μm; the
length of trichomes’ hook was 10.0 ± 1.3 μm.
Occasionally the surface of the main epidermal
cells was covered with waxy structures of
various shapes.
No differences were detected regarding the
control variant on the abaxial leaf surface of
flooded C. canescens plants (Fig. 4 C, D). The
abaxial leaf surface was characterized by the
presence of identical trichomes and zonality
with similar trichomes and elongated stomata.
Localization of silicon inclusions
Control variant. Laser confocal microscopy
allowed us to determine the localization of
silicon inclussions in the leaves of C. canescens.
An evident green fluorescence of silicon
inclussions in the adaxial epidermis was
detected in the control plants (Fig. 5 A).
Silicon inclusions were present in the
form of small bodies of rounded or irregular
shape (at the base of trichomes) and fine-
grained amorphous inclusions smaller than
10 nm (along the trichomes and in the main
epidermal cells). The luminescence intensity
of silicon bodies and amorphous silicon
inclusions differed in the trichomes and main
epidermal cells (Table 1; Fig. 5 C, C’). At the base
Plant Introduction • 91/92 31
Phenotypic plasticity of Corynephorus canescens leaves during flooding
of the trichomes, the fluorescence intensity of
silicon bodies was the highest.
Flooded variant. In flooded plants, the green
fluorescence of silicon inclusions on the
adaxial surface was similar to that in control
samples (Fig. 5 B). Silicon inclusions in the
form of grains were found both at the base of
the trichomes and along their entire cell, as
well as on the surface of the main epidermal
A B
C
D
C’
D’
Figure 5. Microphotographs of silicon inclusions fluorescence in the leaves of control (A, C) and flooded
(B, D) plants of Corynephorus canescens. C’ and D’ – histograms of silicon fluorescence intensity scanned
from the photos C and D (white arrows indicate precise locations); ordinate – silicon fluorescence intensity
(in relative units), abscissa – distance (in μm) scanned from the Figures С and D.
32 Plant Introduction • 91/92
O. Nedukha
cells surrounding the trichomes. The level of
silicon fluorescence in flooded C. canescens
plants is presented in Table 1 and Fig. 5 D, D’.
Comparative analysis of luminescence
intensity showed that flooding increased the
silicon content in trichomes.
Discussion
Thus, C. canescens had the same shape of
leaves (horseshoe-shaped cross-sections)
and their twisting regardless of the growing
conditions. Twisting and/or folding of leaves
is a sign of many psammophytes irrespective
of their place of growth (Ripley & Redmann,
1976; Perrone et al., 2015), which is reflected in
the specialized structure of their leaf lamina
tissues. Change of the leaf shape helps the
plant to reduce water consumption during
increasing external temperature. It optimizes
water and energy balances, responding to
changes in heat capacity, and the conductivity
of water vapor and carbon dioxide (Redmann,
1985).
We found that three weeks of flooding of
C. canescens plants caused a 1.7-time increase
in the leaves’ aerenchyma area compared
to the control. Although the aerenchyma in
flooded plants occupied almost 35 % of the
leaf area, its structure was the same as in
typical plants. Formed aerenchyma provided
more oxygen to the plant and helped in its
resistance against flooding. A similar effect
is reported for many cultivated and wild
plant species in the condition of prolonged
flooding (Vartapetian & Jackson, 1997; Evans,
2004; Parlanti et al., 2011). Therefore, we can
conclude that psammophyte C. canescens can
withstand prolonged flooding.
We found a high density of cuticular-
wax depositions on the periclinal walls of
epidermal cells and the presence of non-
glandular trichomes on the adaxial surface of
C. canescens leaves. Non-glandular trichomes
protect the aboveground plant’s organs
from insects and herbivores, as well as from
mechanical damage (Peter et al., 1995). The
increase in trichomes density also occurs
during drought (Bourland et al., 2003). Dense
pubescence changes the optical properties
of the leaf surface and can reflect or absorb
the light of specific wavelengths. In addition,
trichomes hold an air layer on the leaf surface,
which helps preserve heat and moisture (Peter
et al., 1995).
In epidermal and hypodermal cells of
C. canescens leaves, regardless of growth
conditions, we detected the presence of
inclusions of different shapes and sizes. It is
reported that these inclusions contain calcium
and potassium, and granular inclusions can
be of varying nature (Daniela et al., 2009;
HuaCong et al., 2010). Calcium oxalate crystals,
like silicon inclusions, attract water molecules,
reducing water loss of the plant cells (Zhan-
Yuan, 2000).
The presence of silicon inclusions in
C. canescens cells was proved by laser
confocal microscopy. In C. canescens, silicon
inclusions were found in the main epidermal
cells and trichomes of the adaxial leaf surface,
regardless of growth conditions. According
to the literature, such silicon inclusions
represent a biological form of the silicon ions
connection with proteins, amino acids. lipids
or polysaccharides (Müller & Grachev, 2009).
Silicon ions are often found in combination
with polysaccharides in cell walls, and such
structures, in general, are called silicon
inclusions (Lins et al., 2002; Guerriero et al.,
2016; Grasik et al., 2020). We proved the
presence of silicon ions in such inclusions
using the Pascal program (with laser confocal
microscopy).
Silicon improves the luminous flux
characteristics by keeping the leaf laminas
in a folded state, optimizing photosynthetic
processes (Ma et al., 2011). This also reduces
the heat load due to efficient far-infrared
heat dissipation of silica, which provides a
passive cooling mechanism for folded leaves
during the intense insolation (Wang et al.,
2005). Thus, the obtained data indicate
the phenotypic plasticity of psammophyte
C. canescens realized under the influence of
adverse factors.
Conclusions
The peculiarities of anatomy and surface
ultrastructure of the leaf laminas in
psammophyte C. canescens were revealed. The
localization and content of silicon inclusions
were investigated using laser confocal
microscopy. It was found that the three-
week flooding of C. canescens plants during
Plant Introduction • 91/92 33
Phenotypic plasticity of Corynephorus canescens leaves during flooding
the vegetative growth phase increased the
content of aerenchyma in their leaves and
stimulated the formation of wax depositions
on the adaxial foliar epidermis, and increased
the content of silicone inclusions in foliar
trichomes.
Acknowledgements
We are grateful to Professor D.V. Dubina
(Institute of Botany of NASU), who passed
C. canescens seeds to the Department of
Anatomy and Cell Biology for research. We
are also grateful to Professor, Corresponding
Member of NASU, Elizabeth L. Kordyum
(Institute of Botany of NASU) for reviewing the
manuscript of the article.
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Прояв фенотипової пластичності листків псамофіту Corynephorus canescens при
затопленні
Олена Недуха
Відділ клітинної біології та анатомії, Інститут ботаніки ім. М.Г. Холодного НАН України,
вул. Терещенківська, 2, Київ, 01004, Україна; o.nedukha@hotmail.com
Наведено результати дослідження структури листків псамофіта Corynephorus canescens, що зростав
в контрольованих умовах та при затопленні із використанням методів світлової мікроскопії,
сканувальної електронної мікроскопії та лазерно-конфокальної мікроскопії. Дослідження морфо-
анатомічних показників листків C. canescens у фазі вегетативного росту показало наявність спільних
та відмінних ознак контрольних та затоплених зразків. Спільними ознаками були: форма та розміри
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Plant Introduction • 91/92 35
Phenotypic plasticity of Corynephorus canescens leaves during flooding
пластинок, гіпостоматичний тип пластинки та ізолатеральна структура паренхіми, товстостінний
епідерміс та двошаровий гіподерміс. Відмінними показниками були: наявність ознак руйнації
клітин фотосинтезуючої паренхіми, зміна їхньої форми з формуванням виростів на полюсах
клітин та збільшення майже вдвічі площі аеренхіми у листках C. canescens при затопленні. Методом
сканувальної електронної мікроскопії встановлена подібність ультраструктури та щільності трихом
на адаксіальній поверхні за виключенням формування кутикулярно-воскових структур на поверхні
епідермісу листків у затоплених рослин. Вперше вивчена субклітинна локалізація іонів кремнію
та встановлена наявність аморфних і дрібних кристалічних включень кремнію у периклінальних
стінках основних клітин епідермісу та аморфних включень кремнію – у трихомах листків.
Виявлено підвищення відносного вмісту кремнію вздовж трихоми в епідермісі листків рослин
після затоплення. Припускається, що фенотипічна пластичність C. canescens, що проявлялась у
збільшенні площі аеренхіми в листках та підвищенні вмісту кремнію у трихомах, сприяє оптимізації
як кисневого балансу рослин, так і водного статусу затоплених рослин, і таким чином підвищує
стійкість досліджуваного виду до тривалого затоплення.
Ключові слова: Corynephorus canescens, епідерма листка, затоплення, аеренхіма
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| id | oai:ojs2.plantintroduction.org:article-1593 |
| institution | Plant Introduction |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2025-07-17T12:53:59Z |
| publishDate | 2021 |
| publisher | M.M. Gryshko National Botanical Garden of the NAS of Ukraine |
| record_format | ojs |
| resource_txt_mv | wwwplantintroductionorg/06/5b0080d131377edd7a1fa040edc94c06.pdf |
| spelling | oai:ojs2.plantintroduction.org:article-15932023-08-26T20:39:08Z Appearance of phenotypic plasticity of leaves in psammophyte Corynephorus canescens during flooding Прояв фенотипової пластичності листків псамофіту Corynephorus canescens при затопленні Nedukha, Olena The results of the study of the leaf structure in psammophyte Corynephorus canescens, which grew under controlled conditions and flooding using the methods of light microscopy, scanning electron microscopy, and laser confocal microscopy, are presented. This study revealed common and distinctive signs of morphological and anatomical parameters of C. canescens leaves in the phase of vegetative growth. Among the common features were the shape and size of the leaf laminas, hypostomatic type of the leaf, isolateral structure of the parenchyma, the thick-walled epidermis, and the bilayered hypodermis. Among the distinctive features were the signs of the destruction of cells in the photosynthetic parenchyma, change in their shape with the formation of protuberances at the cells’ poles, and almost doubling area of the aerenchyma in C. canescens leaves under flooding conditions. Scanning electron microscopy showed the similarity of ultrastructure and density of trichomes on the adaxial surface, excepting the formation of cuticular wax structures on the epidermal surface of the leaves in flooded plants. The subcellular localization of silicon inclusions was studied for the first time. The presence of amorphous and small crystalline silicon inclusions in the periclinal walls of the main epidermal cells and amorphous silicon inclusions in leaf trichomes was established. An increase in the relative silicon content along the trichomes in the leaves’ epidermis after flooding was revealed. It was assumed that the phenotypic plasticity of C.&nbsp;canescens, is realized through the increasing area of aerenchyma in leaves and increasing silicon content in trichomes. Such plasticity helps to optimize both the oxygen balance of plants and water balance in flooded plants, thus increasing the species’ resistance to prolonged flooding. Наведено результати дослідження структури листків псамофіта Corynephorus canescens, що зростав в контрольованих умовах та при затопленні із використанням методів світлової мікроскопії, сканувальної електронної мікроскопії та лазерно-конфокальної мікроскопії. Дослідження морфо-анатомічних показників листків C. canescens у фазі вегетативного росту показало наявність спільних та відмінних ознак контрольних та затоплених зразків. Спільними ознаками були: форма та розміри пластинок, гіпостоматичний тип пластинки та ізолатеральна структура паренхіми, товстостінний епідерміс та двошаровий гіподерміс. Відмінними показниками були: наявність ознак руйнації клітин фотосинтезуючої паренхіми, зміна їхньої форми з формуванням виростів на полюсах клітин та збільшення майже вдвічі площі аеренхіми у листках C. canescens при затопленні. Методом сканувальної електронної мікроскопії встановлена подібність ультраструктури та щільності трихом на адаксіальній поверхні за виключенням формування кутикулярно-воскових структур на поверхні епідермісу листків у затоплених рослин. Вперше вивчена субклітинна локалізація іонів кремнію та встановлена наявність аморфних і дрібних кристалічних включень кремнію у периклінальних стінках основних клітин епідермісу та аморфних включень кремнію – у трихомах листків. Виявлено підвищення відносного вмісту кремнію вздовж трихоми в епідермісі листків рослин після затоплення. Припускається, що фенотипічна пластичність C. canescens, що проявлялась у збільшенні площі аеренхіми в листках та підвищенні вмісту кремнію у трихомах, сприяє оптимізації як кисневого балансу рослин, так і водного статусу затоплених рослин, і таким чином підвищує стійкість досліджуваного виду до тривалого затоплення. M.M. Gryshko National Botanical Garden of the NAS of Ukraine 2021-11-21 Article Article application/pdf https://www.plantintroduction.org/index.php/pi/article/view/1593 10.46341/PI2021011 Plant Introduction; No 91/92 (2021); 24-35 Інтродукція Рослин; № 91/92 (2021); 24-35 2663-290X 1605-6574 10.46341/PI91-92 en https://www.plantintroduction.org/index.php/pi/article/view/1593/1518 Copyright (c) 2021 Olena Nedukha http://creativecommons.org/licenses/by/4.0 |
| spellingShingle | Nedukha, Olena Прояв фенотипової пластичності листків псамофіту Corynephorus canescens при затопленні |
| title | Прояв фенотипової пластичності листків псамофіту Corynephorus canescens при затопленні |
| title_alt | Appearance of phenotypic plasticity of leaves in psammophyte Corynephorus canescens during flooding |
| title_full | Прояв фенотипової пластичності листків псамофіту Corynephorus canescens при затопленні |
| title_fullStr | Прояв фенотипової пластичності листків псамофіту Corynephorus canescens при затопленні |
| title_full_unstemmed | Прояв фенотипової пластичності листків псамофіту Corynephorus canescens при затопленні |
| title_short | Прояв фенотипової пластичності листків псамофіту Corynephorus canescens при затопленні |
| title_sort | прояв фенотипової пластичності листків псамофіту corynephorus canescens при затопленні |
| url | https://www.plantintroduction.org/index.php/pi/article/view/1593 |
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