Структурно-функціональні ознаки пластичності листків Typha angustifolia в залежності від умов зростання
The results of the study of leaf anatomy and leaf epidermal ultrastructure of the heliophytic plant Typha angustifolia L. (Typhaceae), which grew in natural conditions: in the water on the bank of the Venetian Strait of the Dnipro River (Kyiv) and on land near the Strait, using light microscopy and...
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M.M. Gryshko National Botanical Garden of the NAS of Ukraine
2025
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| description | The results of the study of leaf anatomy and leaf epidermal ultrastructure of the heliophytic plant Typha angustifolia L. (Typhaceae), which grew in natural conditions: in the water on the bank of the Venetian Strait of the Dnipro River (Kyiv) and on land near the Strait, using light microscopy and scanning electron microscopy are presented. The common and distinctive features of the anatomical signs and the ultrastructure of epidermal cells of T. angustifolia leaves in the phase of vegetative growth of plants were revealed. The anatomical and morphological characteristics of leaves of two ecotypes of T. angustifolia that grew in water and on the terrestrial soil did not differ; the type of mesophyll and the presence of two zones in the epidermis (the zone of cоnvex vault and stomata zone) is stable features for this species. Differences in the size of the leaf blade, the density of stomata, and the density of wax coating on the surface of epidermal cells of the cоnvex vault zone, and also the presence of amorphous silicon in the cell walls of the epidermis are adaptive, and plastic traits that vary depending on the conditions of cattail growth. Besides, scanning electron microscopy of the leaf epidermis of cattail grown in water and on terrestrial soil revealed that growth in water causes the formation of stomata that are deepened into the epidermis, as well as the presence of closed stomata on the lower epidermis, while in the leaves of terrestrial cattail, all stomata were open and located at the same level as the regular epidermal cells. It is assumed that the deepening of stomata into the epidermis contributes to the optimal water balance of leaves under wave action of Strait and high humidity around the leaves of air-water cattail. The obtained results are discussed as a manifestation of phenotypic plasticity and the possible use of epidermal wax as an adaptive marker of heliophytes for growth in different water supply conditions. |
| doi_str_mv | 10.46341/PI2024013 |
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Plant Introduction, 105/106, 3–14 (2025)
© The Authors. This content is provided under CC BY 4.0 license.
RESEARCH ARTICLE
Structural-functional signs of Typha angustifolia leaves plasticity
depending on the growth conditions
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: 30.11.2024 | Accepted: 21.01.2025 | Published online: 16.02.2025
Abstract
The results of the study of leaf anatomy and leaf epidermal ultrastructure of the heliophytic plant
Typha angustifolia L. (Typhaceae), which grew in natural conditions: in the water on the bank of
the Venetian Strait of the Dnipro River (Kyiv) and on land near the Strait, using light microscopy
and scanning electron microscopy are presented. The common and distinctive features of the
anatomical signs and the ultrastructure of epidermal cells of T. angustifolia leaves in the phase of
vegetative growth of plants were revealed. The anatomical and morphological characteristics of
leaves of two ecotypes of T. angustifolia that grew in water and on the terrestrial soil did not differ;
the type of mesophyll and the presence of two zones in the epidermis (the zone of cоnvex vault
and stomata zone) is stable features for this species. Differences in the size of the leaf blade,
the density of stomata, and the density of wax coating on the surface of epidermal cells of the
cоnvex vault zone, and also the presence of amorphous silicon in the cell walls of the epidermis
are adaptive, and plastic traits that vary depending on the conditions of cattail growth. Besides,
scanning electron microscopy of the leaf epidermis of cattail grown in water and on terrestrial
soil revealed that growth in water causes the formation of stomata that are deepened into the
epidermis, as well as the presence of closed stomata on the lower epidermis, while in the leaves
of terrestrial cattail, all stomata were open and located at the same level as the regular epidermal
cells. It is assumed that the deepening of stomata into the epidermis contributes to the optimal
water balance of leaves under wave action of Strait and high humidity around the leaves of air-
water cattail. The obtained results are discussed as a manifestation of phenotypic plasticity and
the possible use of epidermal wax as an adaptive marker of heliophytes for growth in different
water supply conditions.
Keywords: Typha angustifolia, leaves, anatomic signs, epidermis ultrastructure, wax
https://doi.org/10.46341/PI2024013
UDC 581.45 : 58.032 : 549.514.5
Funding: The study was financially supported by the National Academy of Science of Ukraine and performed at the Department of
Cell Biology and Anatomy of the Institute of Botany of the National Academy of Sciences of Ukraine. It was partly supported by the
topic Nr 467 “Cellular and molecular mechanisms of phenotypic plasticity of psammophytes and heliophytes in contrast conditions
of water regime”.
Competing Interests: The author declares that they have no conflict of interest.
4 Plant Introduction • 105/106
Nedukha
Introduction
The problem of plant resistance and
adaptation to adverse environmental changes
not only remains one of the primary problems
of theoretical and experimental biology but is
also significantly exacerbated by the increasing
anthropogenic load on the biosphere and
the forecasts of global climate change, which
threaten to increase air temperature, drought,
and floods. Some of these factors are the
intensity of sunlight, flooding, and changes
in soil moisture, which are characterized by a
combination of changes in the water balance
of plant organs, impaired oxygen respiration
in the root system, and as a result, inhibition
of aerobic processes, impaired absorption of
ions from the soil, lack of nutrients, changes
in metabolic transport, and restrictions on
plant growth and development (Jackson &
Armstrong, 1999; Jackson, 2008; Pennisi,
2008; Malavasi et al., 2016). Despite this,
plants develop mechanisms of adaptation to
unfavorable conditions at different levels of
organization (from morphological to genetic)
that help them to adapt to the changed
environment. It is believed that phenotypic
plasticity is the basis for the survival and
conservation of populations, as well as a
key element of evolution and ecological
relationships of species in habitats (Kordyum,
2012; Dubyna & Kordyum, 2015; Kordyum &
Dubyna, 2019).
A convenient natural model for studying
the adaptive capacity of plants to both natural
flooding and growth under conditions of
varying sunlight and moderate soil drought
may be species that have naturally adapted to
grow in contrasting environmental conditions.
Such a species is the narrow-leave cattail Typha
angustifolia L., a closely related monoecious
wetland plant native to Europe, common in the
temperate zone of the Northern Hemisphere.
Typha angustifolia belongs to the Typhaceae
family; it is a heliophyte, a light-loving
herbaceous perennial aquatic plant (0.8–1.5 m
tall). The leaf arrangement is alternate; the
stem is covered with leaves up to the top.
The stem of the cattail is erect, round, thick,
glabrous, and smooth (Chopyk et al., 1983).
The leaves are linear-lanceolate, flattened. In
aquatic plants, the leaves are straight, while
in terrestrial plants, the leaves can be either
straight or slightly curled inwards.
Because of their widespread distribution
and prevalence, Typha communities form an
important and widespread type of wetland
that provides natural protection against
extreme flooding, conserving freshwater and
improving its quality. They provide habitat for
many species of invertebrates and vertebrates,
providing spawning grounds for fish and
amphibians, and offering birds hiding and
nesting places (Mitsch & Gosselink, 2000;
Bobbink et al., 2006; Heinz, 2010). Typha
species are among the preferred plants in
artificially created wetlands due to their high
carbon dioxide and nutrient uptake capacity
and ability to purify water even in heavy metal
pollution (Saygdeger et al., 2004; Bobbink et al.,
2006). It is known that signs of phenotypic
plasticity of higher plants to changes in soil
moisture include changes in the anatomy of
organs and changes in the ultrastructure of
the leaf epidermis involved in water and gas
transport (Nedukha, 2022). We hypothesized
that similar changes could be observed in
a cattail species growing in contrasting
conditions: water and land. Our study aimed
to investigate the anatomical features of the
leaves of aquatic and dryland cattail plants and
to study the ultrastructure of the leaf blade
surface.
Material and methods
The object of the study was the leaves of the
narrow-leave cattail Typha angustifolia (family
Typhaceae) growing in water on the shore
of the Venetian Gulf (left bank of the Dnipro
River, in the Kyiv area) and on terrestrial soil
about 3–4 m from the shore (Fig. 1). Narrow-
leave cattail plants grew in water at a depth
of 40-50 cm along the shore of the Venetian
Gulf. Terrestrial plants grew on semi-sandy
terrestrial soil near the shore. Material for the
study was collected in the vegetative growth
phase in mid-May. The water temperature
on the collection day was +14 °C, and the air
temperature was +16 °C.
The first leaves from seven plants of each
cattail ecotype were used for microscopic
studies. The material for cytological studies
was fixed in the field. Cuttings from the
broadest part of the leaf blades were
selected for experiments. Leaves that had
completed growth by stretching were used
Plant Introduction • 105/106 5
Structural-functional signs of Typha angustifolia leaves plasticity
for the study. Samples were fixed for light and
electron microscopy with a mixture of 2 %
glutaraldehyde and 2 % paraformaldehyde (1:1,
v/v) in 1 M phosphate buffer, pH 7.2, directly on
the bank, and delivered to the laboratory, where
they were washed with buffer, dehydrated, and
embedded in an epoxy resin mixture (epon–
araldite) according to the generally accepted
method (Bücking & Heyser, 2000). Sections
of 10 μm thickness were made on an RMC
MT-XL ultramicrotome (USA), stained with an
aqueous solution of methyl red, crystal violet-
lactone, and silver amino chromate according
to the protocol (Dayanandan et al., 1983) and
examined under an NF light microscope
(Carl Zeiss, Germany). For scanning electron
microscopy, the fixed material in the aldehyde
mixture was dehydrated and dried according
to the protocol (Bücking & Heyser, 2000).
Then, the leaf blade samples were mounted
on tables, sputtered with gold, and examined
in a scanning electron microscope (JSM 6060
LA) at 30 kV. Statistical processing of stomatal
density and cell size values was performed
using Origin 6.1 software and Student’s t-test
(P ≤ 0.05).
Leaves from seven plants were used to
determine the linear dimensions of the
leaves. Linear cell size was determined on
sections from three water cattail leaves
and three terrestrial cattail leaves. From
each sample, 30–40 epidermal cells and 40-
50 mesophyll cells were taken. A standard
biochemical method based on drying the
samples in a thermostat at + 95 °C until the
weight is constant was used to determine
the relative water content in the leaves. The
obtained cytological and biochemical data
were processed statistically.
To determine the soil moisture content on
which the cattail plants grew, soil samples
were taken at a depth of approximately 20
cm from the surface. The soil samples were
dried in a thermostat at a temperature of
+105 °C to a constant weight. It was found
that the moisture content of the soil on which
the aquatic cattail plants grew was 81 ± 2.3 %,
while the moisture content of the soil on
which the terrestrial cattail plants grew was
57.3 ± 1.9 %.
Results
The leaves of aquatic and terrestrial
T. angustifolia plants were characterized by
a linear-lanceolate shape (Fig. 1), longitudinal
venation, smooth upper surface, and a central
vein distinguished on the lower surface. The
leaves of cattail growing in water were smaller
than those of cattail growing on terrestrial soil.
The average size of the cattail leaves grown
in water was 43.7 ± 5.9 cm along the long axis
and 2.1 ± 0.3 cm along the short axis. Leaves of
terrestrial cattail were 54.3 ± 3.7 cm along the
long axis and 3.2 ± 0.4 cm along the short axis.
The water content in the leaves of water cattail
was 64.7 ± 0.5 %, and in the leaves of terrestrial
cattail – 47.2 ± 0.7 %.
Figure 1. The general appearance of Typha angustifolia plants, which grew in water (A) and terrestrial soil (B).
Comparison of T. angustifolia leaves (C) of the plant grown in water on bank (white arrow) and leaves of
terrestrial plant (black arrows).
B CA
6 Plant Introduction • 105/106
Nedukha
Figure 2. Transverse sections of the middle part of the leaf blades of Typha angustifolia, which grew in the
water (A, C, D, G) and terrestrial plants (B, E, F, H). Scale bars: A, B = 100 µm; C–H = 50 µm.
BA
DC
FE
HG
Plant Introduction • 105/106 7
Structural-functional signs of Typha angustifolia leaves plasticity
Anatomical structure of leaves (Fig. 2)
Leaves of T. angustifolia plants, which grew in
the water
The structure of the leaves is isolateral type
(Fig. 2 A). The leaves were characterized by
the presence of collateral conductive bundles,
which, together with mesophyll cells, divided
the leaf blade into areas in the center of
which a large aerenchyma was formed. The
aerenchyma was formed by the lysis of the
palisade and spongy mesophyll (Fig. 2 C, D).
The average thickness of the leaf blade
was 600 ± 21 μm. The adaxial epidermis is
characterized by rounded cells of different
sizes; the cells surrounding the deepened
stomata were larger than regular epidermal
cells. The height of large epidermal cells was
24.6 ± 2.1 µm, the height of small epidermal cells
was 13 ± 1.2 µm, while the width of large cells
was 25 ± 2.1 µm and small cells was 12.7 ± 1.3 µm,
respectively. The palisade parenchyma cells
were elongated (Fig. 2 C); their size was
50 ± 3.7 × 15 ± 2.1 µm. The average number of
chloroplasts per mesophyll cell was 12.3 ± 1.5.
The cells of the spongy parenchyma varied in
shape from elongated polygonal to rounded
(Fig. 2 D). The size of these cells ranged from
45 ± 3.7 × 21 ± 2.1 µm to 13 ± 1.7 µm (diameter,
rounded cells). The spongy parenchyma cells
surrounding the upper and lower conducting
bundles form a continuous ‘partition’ between
the large air cavities (lysogenic aerenchyma).
The lower epidermis is similar in structure and
cell size to the cells of the upper epidermis
(Fig. 2 D). The height of the large cells of the
lower epidermis was 23.1 ± 2.3 μm, the height
of the small epidermal cells was 12.4 ± 1.2 μm,
while the width of these cells was 25.2 ± 2.1 μm
and 12.1 ± 1.3 μm, respectively.
Leaves of terrestrial T. angustifolia plants
The anatomical and morphological
characteristics of terrestrial cattail plants
were similar to those that grew in water; the
structure of the leaf blade was also isolateral
(Fig. 2 B, E, F). The average thickness of the
leaf blade was 623 ± 32 µm. The shape of the
epidermal and mesophyll cells was similar to
that of the leaves of air-water cattail plants.
The average size of large cells (height × width)
of the upper epidermis was 23.3 ± 2.2 × 24 ± 2.5
µm, the size of small cells was 12.3 ± 1.2 ×
13 ± 1.9 µm; of the lower epidermis – 21.3 ± 2.1
× 20 ± 2. 1 µm, and small cells – 11.3 ± 1.2 ×
12 ± 1.9 µm; palisade mesophyll cells –51 ± 3.3
× 13 ± 2.0 µm and spongy mesophyll cells –
from 42 ± 3.7 × 21 ± 0.9 to 10 ± 1.3 (for rounded
cells) µm, respectively. The average number of
chloroplasts per section of mesophyll cells was
13.4 ± 2.5.
It should be noted that the mesophyll cells
and cells of the conducting bundles in the
leaves of two cattail ecotypes on the sections
were blue stained. In contrast, epidermal
cells, especially of the adaxial surface around
the perimeter of all epidermal cell walls, were
stained in bright yellow hot color with a red
tint, which is visible at high magnification of
the microscope after staining the sections with
an aqueous solution of methyl red, crystalline
violet lactone and silver amino chromate
(Fig. 2 G, H). Similar staining could be observed
occasionally in the palisade parenchyma cell
walls in contact with the upper epidermis in
the leaves of terrestrial cattail (Fig. 2 E).
Microstructure of T. angustifolia leaves
epidermal surface (Figs. 3, 4)
The study of the ultrastructure of the surface
of leaves of aquatic and terrestrial plants of
narrow-leave cattail revealed the presence of
two zones of different structures on the adaxial
and abaxial surfaces: the cоnvex vault zone and
the stomatal zone (Figs. 3, 4). Differences in
the ultrastructure of the surface of air-water
and terrestrial plants were manifested in the
density of stomata, the presence and structure
of waxy inclusions on the cell surface, as well
as in the deepening of stomata in the epidermis
of water cattail leaves.
The leaf epidermis of T. angustifolia plants,
which grew in the water (Fig. 3)
Adaxial surface (Fig. 3 A–C). The width of the
cоnvex vault zone ranged from 22 ± 2.1 to 53 ± 4.1
µm; the width of the stomata zone ranged
from 68 ± 3.7 to 82 ± 7.4 µm, respectively. On the
surface of the cоnvex vault zone of the upper
epidermis, individual cells were distinguished,
the anticlinal walls of which protruded above
the periclinal walls, and the thickness of the
anticlinal walls was ca. 5 µm. Trichomes and
waxy inclusions were absent on the surface
of epidermal cells of the cоnvex vault zone.
Meanwhile, in the stomatal zone, the surface
of ordinary epidermal cells was covered with
waxy structures of various shapes (Fig. 3 B, C):
8 Plant Introduction • 105/106
Nedukha
needle-shaped, drop-shaped, or triangular.
Wax inclusions were of different sizes (1 to 3
µm along the long axis). Stomatal density was
534 ± 17 per mm2 area. Stomata are elongated
and deepened. Epidermal cells located around
the stomata are raised. The surface of the
stomatal guard cells is smooth, without wax.
Stomata size was 18 ± 1.1 µm along the long axis
and 14 ± 1.7 µm along the short axis.
Abaxial surface (Fig. 3 D–F). It is also
characterized by the presence of two
structural zones, similar to those on the
upper surface of the leaf in terms of the
ultrastructure of epidermal cells and the
BA
DC
FE
1
2
1
2
Figure 3. The ultrastructure of the adaxial (A–C) and abaxial (D–F) leaf surfaces of Typha angustifolia plants,
which grew in the water. The black curved lines (A & D) indicate: 1 – cоnvex vault zone; 2 – stomata zone.
Closed stomata are indicated by black arrows (E).
Plant Introduction • 105/106 9
Structural-functional signs of Typha angustifolia leaves plasticity
1
1
2
2
BA
DC
FE
Figure 4. The ultrastructure of the adaxial (A–C) and abaxial (D–F) leaf surfaces of terrestrial Typha
angustifolia plants. The black curved lines (A & D) indicate: 1 – cоnvex vault zone; 2 – stomata zone.
structure of stomatal guard cells. The only
difference was a decrease in the width of the
cоnvex vault zone to 21 ± 2.0 µm. The density
of stomata on the lower epidermis was 610 ± 23
per mm2; closed stomata were observed along
with open stomata (Fig. 3 E; lower row of
closed stomata cells indicated by arrows).
The leaf epidermis of terrestrial T. angustifolia
plants (Fig. 4)
Adaxial surface. The presence of two
structurally distinct zones is also characteristic
of terrestrial cattail leaves (Fig. 4). The surface
of the cоnvex vault zone of the epidermis is
covered with wax structures. The width of
10 Plant Introduction • 105/106
Nedukha
the cоnvex vault zone varies from 45.2 ± 4.1
to 55.0 ± 5.7 µm. On the surface of the cоnvex
vault, anticlinal walls are visible, similar to
those of water cattail, with a width of 4 to 6
µm. The anticlinal protruding and recessed
periclinal walls are densely covered with wax
structures (plaque) of various shapes: round,
triangular, or square. Wax structures can also
be seen on the periphery of stomatal guard
cells (Fig. 4 C). The stomata are elongated, with
a density of 414 ± 12 per mm2 area.
Abaxial surface. Abaxial epidermis is
characterized by the presence of two zones
similar to those on the upper epidermis: the
cоnvex vault zone and stomata zones (Fig. 4
D–F). The average width of the stomata zone
ranges from 50 ± 3.7 to 67 ± 7.1 µm, and the width
of the cоnvex vault zone is from 23 ± 3.7 to
50 ± 4.1 µm, with arrow-shaped (or triangular)
wax structures clearly visible on the surface of
the tubercle zone. The surfaces of the stomatal
guard cells, which are not deepened but are
almost at the level of the regular epidermal
cells, are encrusted with wax structures (Fig. 4
E, F). The density of stomata on the lower
epidermis is 458 ± 23 per mm2 of area.
Discussion
Thus, our studies of the anatomical and
morphological characteristics of the leaves of
aquatic and terrestrial cattail T. angustifolia
showed that, regardless of the place of growth
of cattail, the leaves were characterized
by a similar anatomical structure with an
amphistomatic type of structure and large
aerenchyma cavities. A similar anatomical
structure of leaves is also characteristic for
other Typhaceae, including T. latifolia L.
growing in Eurasia, namely, an amphistomatic
type of leaf blades with sclerenchymal
conductive bundles that connect palisade
and spongy parenchyma, forming partitions
between large aerenchymal formations
(Henry, 2003).
Despite the identity of the anatomical
structure of the studied objects, we found
differences in the size of leaf blades of aquatic
and terrestrial cattail plants: significantly
larger leaf sizes of terrestrial cattails than
those in plants grown in the water. There
may be several reasons for the differences in
leaf size: lack of nutrients for water cattail,
different cell cycle rates, different ploidy,
and differences in population density. The
growth of higher plants in water is known to
be characterized by both partial hypoxia and a
particular nutrient limitation, which is typical
for different grass populations (Insausti et al.,
2001). We also do not exclude the possibility
that the aquatic and terrestrial cattail plants
that we studied are characterized by different
ploidy, which is known to affect morphological
traits; in particular, such a relationship has
been established for reed leaves (Clevering &
Lissner, 1999; Paucă-Comănescu et al., 1999;
Hansen et al., 2007). Researchers have shown
that the lower the ploidy, the smaller the cell
size (Hansen et al., 2007). This relationship
between ploidy level and morphology is
generally biological, as a relevant and general
effect of polyploidy is increased cell size.
However, polyploidy does not always lead to an
overall increase in plant size, as a general effect
of polyploidy can also be a decrease in the
number of cell divisions during development
(Stebbins, 1971).
Particularly noteworthy are the data
of researchers who studied the effect of
T. angustifolia and T. domingensis Pers.
population density on the anatomical and
physiological characteristics of these species
(Corrêa et al., 2015). The authors showed
that an increased growth rate characterizes
plants from populations with high cattail
density. Corrêa et al. (2015) attribute this
phenomenon to lower apoplastic barriers in
the roots and to an increased ability to absorb
nutrients, as well as to a particular root/
shoot size ratio, compared to plants with
low density (less than 50 % of the colonizing
capacity). Given these data, we assume that
the morphological structure of the leaves of
the two cattail populations we studied was
also influenced by the density of these plants:
the land-based population with high density
had larger leaf sizes than the cattail plants
growing at low density (in water). Thus, the
complex structural, physiological, and genetic
mechanisms of leaf plasticity of narrow-leave
cattail plants growing in the Kyiv area require
further investigation.
In addition to the differences mentioned
above, we also found certain differences in
the stain of the cell walls of mesophyll and
epidermal tissues of the studied samples of
narrow-leave cattail. The walls of epidermal
Plant Introduction • 105/106 11
Structural-functional signs of Typha angustifolia leaves plasticity
tissues, especially the adaxial epidermis,
acquired a hot yellow color with a red tint,
regardless of the place of growth of the
species. The mesophyll walls adjacent to
the adaxial epidermal cells in the leaves of
the terrestrial cattail also had a similar color
(Fig. 2 E, H), while the mesophyll walls of the
leaves of aquatic cattail plants were blue. The
peculiarity of this phenomenon is the physical
property of the dye – methyl red, a crystalline
violet lactone, which, when combined with
silicon ions (SiOH), dyes amorphous silicon in
cells in such bright color (Dayanandan et al.,
1983; Guerriero et al., 2020). Based on the
above data, we assume that the epidermal
walls of cattail leaves contain amorphous
silicon, similar to those in the leaves of many
plant species, including leaves of rice, bamboo,
and other species (Dayanandan et al., 1983;
Blecher et al., 2012; Guerriero et al., 2020).
Obviously, the presence of amorphous silicon
in the walls of the epidermis and mesophyll of
narrow-leave cattail leaves is a structural and
functional feature that determines the optimal
water and osmotic balance of the leaves.
Our data on the ultrastructure of the leaf
surface of aquatic and terrestrial cattails
revealed both common and different features.
The common feature was the presence of two
structurally different zones (stomatal and
cоnvex vault) in the epidermis. Whereas the
differences were manifested in the density of
stomata, the increase of wax coating on the
surface of epidermal cells in the cоnvex vault
zone and the stomatal zone, as well as the
presence of wax on the periphery of stomatal
guard cells in the leaves of dry cattail. The
presence of two zones in the leaf epidermis
is also characteristic of other species of this
family, in particular, T. domingensis (Cruz et al.,
2019) and T. angustifolia, which grew in Brazil
(Corrêa et al., 2015).
We also discovered a phenomenon -
almost half of the stomata on the lower
epidermis of the water cattail were closed
entirely. These data require an explanation.
On the one hand, it is known that stomatal
functions are associated not only with the
regulation of water balance (transpiration
and cell osmotic pressure), but also with CO2
uptake for photosynthesis (Boyer et al., 1997;
Lawson & Blatt, 2014; Roche, 2015). Hence,
stomatal conductance is a key physiological
parameter that regulates plant growth
and development under normal and stress
conditions (Hetherington & Woodward, 2003;
Hasanuzzaman et al., 2023). Aquatic cattail
plants with higher stomatal conductance (on
the upper epidermis) are characterized by a
high rate of CO2 assimilation. The presence
of a large proportion of closed stomata on
the lower epidermis of aquatic cattail leaves
can be explained by two reasons. First,
some stomata are temporarily inoperative,
as natural conditions can induce surface-
specific stomatal closure (Richardson et al.,
2017). The second reason is the complete non-
functionality of a part of stomata in water
cattail, similar to that described for the leaves
of aquatic plants, in particular Salvinia herzogii
de la Sota, Lemna minor L. (Ziegler, 1987; de la
Sota et al., 1990; Shtein et al., 2017), Nymphaea
violacea Lehm. and others (Kaul, 1976; Ziegler,
1987), in which gas exchange occurs only
through the upper surface of the leaves. In
addition, the closure of stomata on the lower
epidermis of water cattail may be associated
with the fact that the lower surface can be
affected by the water environment and waves,
and this surface can also be in contact with the
surrounding aquatic microflora and numerous
algae (Tyree, & Cheung, 1977; Nedukha, 2011).
When studying the ultrastructure of the
epidermis of cattail leaves, we found an
increase in the content and density of wax
inclusions on the surface of the epidermal
walls. It is known that the wax formed on
the outer surface of epidermal cells inhibits
transpiration and can absorb and/or reflect
sunlight (Kolattukudy, 1981; Schönherr, 1982;
Kerstiens, 1996, 2006). The wax formed on
the cell surface, together with cuticular wax,
is one of the barriers to water evaporation
and an inhibitor of the transport of small
water molecules both from the epidermis
and their transport into the cells (Hauke &
Schreiber, 1998; Barthlott et al., 2017). Based
on this literature and our scanning electron
microscopy data, we can assume that the ‘exit’
of cattail plants from water to terrestrial soil
(to land) causes an increase in the formation
of wax in the leaf epidermis, and this is a
manifestation of the phenotypic plasticity
of the plants of the studied species. We
believe that wax structures on the surface of
epidermal cells can be a marker of changes (or
decreases) in water transport in plant leaves.
Using these traits in breeding and/or genetic
12 Plant Introduction • 105/106
Nedukha
studies will allow researchers to select species
that can adapt to environmental stresses, such
as drought or changes in plant water supply.
Conclusions
The anatomical and morphological
characteristics of leaves of two ecotypes of
Typha angustifolia grown in water and on
the terrestrial soil did not differ: the type of
mesophyll and the presence of two zones
in the epidermis: the zone of cоnvex vault
and stomata zone is stable features for this
species. Differences in the density of stomata
and the density of wax coating on the surface
of epidermal cells indicated the phenotypic
plasticity of the species and the modification
of leaf structure depending on the growth
conditions, particularly soil moisture. Scanning
electron microscopy of the leaf epidermis
of cattail grown in water and on terrestrial
soil revealed that growth in the water causes
the formation of stomata that are deepened
into the epidermis, as well as the presence of
closed stomata on the lower epidermis, while
in the leaves of terrestrial cattail, all stomata
were open and located at the same level as
the regular epidermal cells. It is assumed that
the deepening of stomata into the epidermis
contributes to the optimal water balance of
leaves under conditions of wave action and
high humidity around the leaves of the water
cattail.
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14 Plant Introduction • 105/106
Nedukha
Структурно-функціональні ознаки пластичності листків Typha angustifolia в
залежності від умов зростання
Олена Недуха
Відділ клітинної біології та анатомії, Інститут ботаніки імені М.Г. Холодного НАН України,
вул. Терещенківська, 2, Київ, 01601, Україна; o.nedukha@hotmail.com
Наведено результати дослідження анатомії та ультраструктури епідермісу листків геліофітної
рослини Typha angustifolia L. (Typhaceae), яка зростала в природних умовах: у воді на березі
Венеціанської протоки р. Дніпро (м. Київ) та на суші поблизу берега протоки, методами світлової
мікроскопії та скануючої електронної мікроскопії. Виявлено спільні та відмінні риси анатомічних
ознак та ультраструктури клітин епідермісу листків T. angustifolia у фазі вегетативного росту рослин.
Анатомо-морфологічні ознаки листків двох екотипів T. angustifolia, що зростали у воді та на суходолі,
не відрізнялися; тип мезофілу та наявність двох зон в епідермісі (зона опуклого склепіння та зона
продихів) були стабільними ознаками для цього виду. Відмінності в розмірах листкової пластинки,
щільності продихів та щільності воскового нальоту на поверхні клітин епідермісу зони випуклого
склепіння, а також наявність аморфного кремнію в клітинних стінках епідермісу є адаптивно-
пластичними ознаками, які змінюються залежно від умов зростання рогозу. Крім того, скануюча
електронна мікроскопія епідермісу листків рогозу, вирощеного у воді та на суходолі, показала,
що зростання рослини у воді спричиняє формування продихів, заглиблених в епідерміс, а також
наявність закритих продихів на нижньому епідермісі, тоді як у листках наземного рогозу всі
продихи відкриті і розташовані на одному рівні з основними клітинами епідермісу. Припускається,
що заглиблення продихів в епідерміс сприяє оптимальному водному балансу листків в умовах
хвильової дії затоки та підвищеної вологості навколо листків повітряно-водного рогозу. Отримані
результати обговорюються як прояв не тільки фенотипової пластичності, але й можливого
використання епідермального воску як адаптивного маркера геліофітів для зростання в різних
умовах водозабезпечення.
Ключові слова: Typha angustifolia, листки, анатомічні ознаки, ультраструктура епідермісу, віск
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| id | oai:ojs2.plantintroduction.org:article-1646 |
| institution | Plant Introduction |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2025-07-28T04:16:14Z |
| publishDate | 2025 |
| publisher | M.M. Gryshko National Botanical Garden of the NAS of Ukraine |
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| resource_txt_mv | wwwplantintroductionorg/70/298eb572dc57b6d9d02ad894a3239d70.pdf |
| spelling | oai:ojs2.plantintroduction.org:article-16462025-07-27T21:03:35Z Structural-functional signs of Typha angustifolia leaves plasticity depending on the growth conditions Структурно-функціональні ознаки пластичності листків Typha angustifolia в залежності від умов зростання Nedukha, Olena The results of the study of leaf anatomy and leaf epidermal ultrastructure of the heliophytic plant Typha angustifolia L. (Typhaceae), which grew in natural conditions: in the water on the bank of the Venetian Strait of the Dnipro River (Kyiv) and on land near the Strait, using light microscopy and scanning electron microscopy are presented. The common and distinctive features of the anatomical signs and the ultrastructure of epidermal cells of T. angustifolia leaves in the phase of vegetative growth of plants were revealed. The anatomical and morphological characteristics of leaves of two ecotypes of T. angustifolia that grew in water and on the terrestrial soil did not differ; the type of mesophyll and the presence of two zones in the epidermis (the zone of cоnvex vault and stomata zone) is stable features for this species. Differences in the size of the leaf blade, the density of stomata, and the density of wax coating on the surface of epidermal cells of the cоnvex vault zone, and also the presence of amorphous silicon in the cell walls of the epidermis are adaptive, and plastic traits that vary depending on the conditions of cattail growth. Besides, scanning electron microscopy of the leaf epidermis of cattail grown in water and on terrestrial soil revealed that growth in water causes the formation of stomata that are deepened into the epidermis, as well as the presence of closed stomata on the lower epidermis, while in the leaves of terrestrial cattail, all stomata were open and located at the same level as the regular epidermal cells. It is assumed that the deepening of stomata into the epidermis contributes to the optimal water balance of leaves under wave action of Strait and high humidity around the leaves of air-water cattail. The obtained results are discussed as a manifestation of phenotypic plasticity and the possible use of epidermal wax as an adaptive marker of heliophytes for growth in different water supply conditions. Наведено результати дослідження анатомії та ультраструктури епідермісу листків геліофітної рослини Typha angustifolia L. (Typhaceae), яка зростала в природних умовах: у воді на березі Венеціанської протоки р. Дніпро (м. Київ) та на суші поблизу берега протоки, методами світлової мікроскопії та скануючої електронної мікроскопії. Виявлено спільні та відмінні риси анатомічних ознак та ультраструктури клітин епідермісу листків T. angustifolia у фазі вегетативного росту рослин. Анатомо-морфологічні ознаки листків двох екотипів T. angustifolia, що зростали у воді та на суходолі, не відрізнялися; тип мезофілу та наявність двох зон в епідермісі (зона опуклого склепіння та зона продихів) були стабільними ознаками для цього виду. Відмінності в розмірах листкової пластинки, щільності продихів та щільності воскового нальоту на поверхні клітин епідермісу зони випуклого склепіння, а також наявність аморфного кремнію в клітинних стінках епідермісу є адаптивно-пластичними ознаками, які змінюються залежно від умов зростання рогозу. Крім того, скануюча електронна мікроскопія епідермісу листків рогозу, вирощеного у воді та на суходолі, показала, що зростання рослини у воді спричиняє формування продихів, заглиблених в епідерміс, а також наявність закритих продихів на нижньому епідермісі, тоді як у листках наземного рогозу всі продихи відкриті і розташовані на одному рівні з основними клітинами епідермісу. Припускається, що заглиблення продихів в епідерміс сприяє оптимальному водному балансу листків в умовах хвильової дії затоки та підвищеної вологості навколо листків повітряно-водного рогозу. Отримані результати обговорюються як прояв не тільки фенотипової пластичності, але й можливого використання епідермального воску як адаптивного маркера геліофітів для зростання в різних умовах водозабезпечення. M.M. Gryshko National Botanical Garden of the NAS of Ukraine 2025-02-16 Article Article application/pdf https://www.plantintroduction.org/index.php/pi/article/view/1646 10.46341/PI2024013 Plant Introduction; No 105/106 (2025); 3-14 Інтродукція Рослин; № 105/106 (2025); 3-14 2663-290X 1605-6574 10.46341/PI105-106 en https://www.plantintroduction.org/index.php/pi/article/view/1646/1561 Copyright (c) 2025 Olena Nedukha http://creativecommons.org/licenses/by/4.0 |
| spellingShingle | Nedukha, Olena Структурно-функціональні ознаки пластичності листків Typha angustifolia в залежності від умов зростання |
| title | Структурно-функціональні ознаки пластичності листків Typha angustifolia в залежності від умов зростання |
| title_alt | Structural-functional signs of Typha angustifolia leaves plasticity depending on the growth conditions |
| title_full | Структурно-функціональні ознаки пластичності листків Typha angustifolia в залежності від умов зростання |
| title_fullStr | Структурно-функціональні ознаки пластичності листків Typha angustifolia в залежності від умов зростання |
| title_full_unstemmed | Структурно-функціональні ознаки пластичності листків Typha angustifolia в залежності від умов зростання |
| title_short | Структурно-функціональні ознаки пластичності листків Typha angustifolia в залежності від умов зростання |
| title_sort | структурно-функціональні ознаки пластичності листків typha angustifolia в залежності від умов зростання |
| url | https://www.plantintroduction.org/index.php/pi/article/view/1646 |
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