Морфологічні ознаки листків ломиносів як відображення їх екологічних особливостей
Quantitative morpho-anatomical features of leaves of nine Clematis taxa (C. alpina ‘Pamela Jackman’, C. macropetala ‘Maidwell Hall’, C. integrifolia ‘Aljonushka’, C. ispahanica ‘Zvezdograd’, C. fargesii ‘Paul Farges’, C. texensis ‘Princess Diana’, C. tibetana, C. viticella,...
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| Дата: | 2021 |
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M.M. Gryshko National Botanical Garden of the NAS of Ukraine
2021
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Plant Introduction| _version_ | 1860145089561493504 |
|---|---|
| author | Kovalyshyn, Iryna Pinchuk, Andrii Likhanov, Artur |
| author_facet | Kovalyshyn, Iryna Pinchuk, Andrii Likhanov, Artur |
| author_sort | Kovalyshyn, Iryna |
| baseUrl_str | https://www.plantintroduction.org/index.php/pi/oai |
| collection | OJS |
| datestamp_date | 2023-08-26T20:39:20Z |
| description | Quantitative morpho-anatomical features of leaves of nine Clematis taxa (C. alpina ‘Pamela Jackman’, C. macropetala ‘Maidwell Hall’, C. integrifolia ‘Aljonushka’, C. ispahanica ‘Zvezdograd’, C. fargesii ‘Paul Farges’, C. texensis ‘Princess Diana’, C. tibetana, C. viticella, and C. heracleifolia) were determined with the aim to analyze their adaptation to the environmental conditions.Among investigated clematises, there were plants with hypostomatic (C. viticella, C. fargesii ‘Paul Farges’, C. heracleifolia, C. texensis ‘Princess Diana’, C. macropetala ‘Maidwell Hall’, and C. alpina ‘Pamela Jackman’), and amphistomatic leaves (C. ispahanica ‘Zvezdograd’ and C. tibetana). In C. integrifolia ‘Aljonushka’ leaves were hypostomatic, but few solitary stomata were also present on the adaxial surface. In the leaves of investigated taxa, the palisade coefficient ranged from 27.3% (C. alpina ‘Pamela Jackman’) to 49.9% (C. tibetana). The leaves also differed significantly in size. In particular, leaves of C. integrifolia ‘Aljonushka’ were almost ten times smaller than such of C. heracleifolia.As a result of UPGMA clustering, the plants that can survive in severe windy weather in open rocky areas, Clematis tibetana and C. ispahanica ‘Zvezdograd’, were joined in a separate cluster. The second cluster combined C. alpina ‘Pamela Jackman’ and C. macropetala ‘Maidwell Hall’ – cultivars blooming in the spring, during a period of significant difference in daily temperatures. A relatively small leaf area in plants from these two clusters may indicate an adaptation by reducing the transpiration area and general windage. The third cluster united the rest of investigated taxa, mostly – the mesophytic plants with a relatively large leaf area. However, due to similar morpho-anatomical structure of the leaf, the third cluster also comprised C. integrifolia ‘Aljonushka’ with the smallest leaves. |
| doi_str_mv | 10.46341/PI2021002 |
| first_indexed | 2025-07-17T12:53:42Z |
| format | Article |
| fulltext |
Plant Introduction, 89/90, 89–100 (2021)
© The Authors. This content is provided under CC BY 4.0 license.
RESEARCH ARTICLE
Morphological features of the leaves in clematises as a reflection of their
ecological peculiarities
Iryna Kovalyshyn 1, *, Andrii Pinchuk 2, Artur Likhanov 2
1 Institute of Plant Physiology and Genetics, National Academy of Science of Ukraine, Vasylkivska str. 31/17, 03022 Kyiv, Ukraine;
* Iryna_B_Kovalyshyn@ukr.net
2 National University of Life and Environmental Sciences of Ukraine, Heroyiv Oborony str. 15, 03041 Kyiv, Ukraine
Received: 13.01.2021 | Accepted: 01.06.2021 | Published online: 14.06.2021
Abstract
Quantitative morpho-anatomical features of leaves of nine Clematis taxa (C. alpina ‘Pamela Jackman’,
C. macropetala ‘Maidwell Hall’, C. integrifolia ‘Aljonushka’, C. ispahanica ‘Zvezdograd’, C. fargesii ‘Paul Farges’,
C. texensis ‘Princess Diana’, C. tibetana, C. viticella, and C. heracleifolia) were determined with the aim to analyze
their adaptation to the environmental conditions.
Among investigated clematises, there were plants with hypostomatic (C. viticella, C. fargesii ‘Paul Farges’,
C. heracleifolia, C. texensis ‘Princess Diana’, C. macropetala ‘Maidwell Hall’, and C. alpina ‘Pamela Jackman’),
and amphistomatic leaves (C. ispahanica ‘Zvezdograd’ and C. tibetana). In C. integrifolia ‘Aljonushka’ leaves
were hypostomatic, but few solitary stomata were also present on the adaxial surface. In the leaves
of investigated taxa, the palisade coefficient ranged from 27.3% (C. alpina ‘Pamela Jackman’) to 49.9%
(C. tibetana). The leaves also differed significantly in size. In particular, leaves of C. integrifolia ‘Aljonushka’
were almost ten times smaller than such of C. heracleifolia.
As a result of UPGMA clustering, the plants that can survive in severe windy weather in open rocky areas,
Clematis tibetana and C. ispahanica ‘Zvezdograd’, were joined in a separate cluster. The second cluster
combined C. alpina ‘Pamela Jackman’ and C. macropetala ‘Maidwell Hall’ – cultivars blooming in the spring,
during a period of significant difference in daily temperatures. A relatively small leaf area in plants from
these two clusters may indicate an adaptation by reducing the transpiration area and general windage.
The third cluster united the rest of investigated taxa, mostly – the mesophytic plants with a relatively
large leaf area. However, due to similar morpho-anatomical structure of the leaf, the third cluster also
comprised C. integrifolia ‘Aljonushka’ with the smallest leaves.
Keywords: Clematis, cluster analysis, leaf, epidermis, mesophyll
https://doi.org/10.46341/PI2021002
UDC 581.4 : 582.65
Authors’ contributions: Iryna Kovalyshyn collected and contributed data, performed the analysis, wrote the paper. Andrii Pinchuk
conceived and designed the analysis, wrote the paper. Artur Likhanov conceived and designed the analysis, performed the analysis.
Funding: The research was carried out at the expense of state funding.
Competing Interests: The authors declare no conflict of interest.
Introduction
The genus Clematis L. (Ranunculaceae Juss.)
unites valuable ornamental and medicinal
plants of various life forms, which grow on
all continents except Antarctica (Sneddon,
1975; Takhtajan, 1978; Tamura, 1993; Moon
et al., 2013; Zhang et al., 2013). Due to the
https://creativecommons.org/licenses/by/4.0/
https://orcid.org/0000-0002-5280-6693
https://orcid.org/0000-0003-1256-9838
https://orcid.org/0000-0001-6580-7241
90 Plant Introduction • 89/90
I. Kovalyshyn, A. Pinchuk, A. Likhanov
long history of introduction and selection,
many clematises are available for planting
in a moderate climatic zone and are
successfully cultivated in Ukrainian botanical
gardens (Rijekstynja & Rijekstynysh, 1990;
Tamura, 1993; Vachnovskaya, 2007). Taking
into account the wide range of decorative
application, clematises are good plants for
urban gardening.
Clematises differ in the rhythms of their
development and can be deciduous or
evergreen. They are mainly entomophilous,
monoecious or dioecious plants with
opposite pinnate leaves. Morpho-anatomical
diversity of their leaves is often determined
by the ecological conditions of their habitats
(they often occupy forest edges, rocky
slopes, rocky river banks, and also steppes
and desert landscapes) in different climatic
zones (Rijekstynja & Rijekstynysh, 1990;
Tamura, 1993; Buisson & Lee, 1993; Bååth &
Anderson, 2003; Ahmad et al., 2016; Arimy
et al., 2017).
Adaptation to environmental conditions is
reflected in the size, orientation, and density
of structural elements of the leaf (Pashkevych,
2014; Arimy et al., 2017; Zanão Júnior et al.,
2017; Araujo et al., 2019; Ornellas et al., 2019).
The mechanisms of such adaptation may differ
(e.g., by changing the area and thickness of
the leaf blade, mesophyll or epidermal cell
sizes, number of stomata, or developing the
pubescence), but some genetically determined
traits remain intact (e.g., the type of leaf
blade, vascularization, stomatal apparatus,
and trichomes). Therefore, the anatomical
structure of the leaves often serves as a
valuable source of taxonomic features (Isnard
et al., 2003; Jinghua & Liangqian, 2003; Javed
et al., 2012). Taking into account these two
aspects, we analyzed morpho-anatomical
features of the leaves of nine Clematis taxa
to evaluate their adaptive potential to urban
conditions.
Material and methods
Morphological peculiarities were analyzed
for nine Clematis taxa (Table 1), which were
Taxon Breeding technology Natural range
C. alpina (L.) Mill.
‘Pamela Jackman’
Selection from C. alpina seed
offsprings
Forests in the mountains of Central and Eastern
Europe (rocky slopes, river banks and bushes)
C. macropetala Ladeb.
‘Maidwell Hall’
Selection from C. macropetala seed
offsprings
Open shaded slopes, edges, coniferous and
deciduous forests of Eastern Siberia, the Far East,
China, North Korea, and Eastern Mongolia
C. integrifolia L.
‘Aljonushka’
Hybridization C. jackmanii
‘Nezhdannyi’ × C. іntegrifolia
Clematis integrifolia is common among shrubs,
on the edges and banks of rivers of Crimea,
Prykarpattia, Western Europe, the Caucasus,
Northern Kazakhstan, Turkey, Western China.
Clematis jackmanii `Nezhdannyì was obtained by
free pollination of C. jackmanii
C. ispahanica Boiss.
‘Zvezdograd’
Mutagenesis of C. ispahanica Clematis ispahanica grows on steppe slopes in
Central Asia and Iran
C. fargesii Franch.
‘Paul Farges’
Hybridization C. fargesii ×
C. vitalba L.
Clematis fargesii grows in West China. Clematis
vitalba occurs in Crimea, Caucasus, Central and
Southern Europe, Asia Minor, and North Africa
C. texensis Buckl.
‘Princess Diana’
Hybridization Clematis ‘Bees’ Jubilee’
× C. texensis
Clematis texensis grows in USA (Texas). Origin of
Clematis ‘Bees’ Jubilee’ is unknown
C. tibetana Kuntze Southeast Asia (Indian Himalayas, Nepal, China, and
Tibet)
C. viticella L. Rocky slopes and edges in Southern Europe,
Western Caucasus, Iran, Asia Minor
C. heracleifolia DC. East China and Korea
Table 1. Investigated Clematis taxa and their natural ranges following Beskaravaynaya (1998) and Clematis
on the Web (2021).
Plant Introduction • 89/90 91
Morphological features of the leaves in clematises and their ecological peculiarities
chosen due to their high ornamental features
combined with the possibility of growing
in public urban plantations. Selected plants
vary phenologically and morphologically,
so they can be applied for the creation of
specific landscape compositions. They also
differ by origin, which causes some ecological
preferences and peculiarities.
For the morphological analysis, for each
taxon five mature leaves of the brunch
middle part were selected from the western
side of plant habitus. Middle parts of leaflets
from selected leaves were used for cuts and
preparation of the permanent slides. Imprints
of abaxial and adaxial sides of mature leaves
were applied to determine the structure of
the epidermis (Evert, 2005). The anatomical
structure of the leaves was studied using
permanent (10 μm thick) and native slides.
Differentiated staining of the leaf tissues was
performed using safranin (stains lignified
elements) and water blue (stains unlignified
elements) dyes (Furst, 1979).
The palisade coefficient indicates the
percentage of palisade mesophyll layer from
total mesophyll layer in the leaf blade and was
calculated by the formula:
K = 100 * MP / MP+S, where
K – the palisade coefficient, in %,
MP – the height of the palisade mesophyll
cells layer on the cross-section of the leaf, in
µm
MP+S – the total height of the mesophyll, in
µm.
Following leaf characters were analysed:
1 – leaf blade thickness, μm; 2 – height of
the upper epidermis cells, μm; 3 – width
of upper epidermis cell, μm; 4 – height
of lower epidermis cells, μm; 5 – width of
lower epidermis cells, μm; 6 – thickness of
palisade mesophyll layer, μm; 7 – height of
palisade mesophyll cell, μm; 8 – width of
palisade mesophyll cell, μm; 9 – thickness of
spongy mesophyll, μm; 10 – height of spongy
mesophyll cells, μm; 11 – width of spongy
mesophyll cells, μm; 12 – diameter of vascular
bundle with sheath cells, μm; 13 – diameter of
vascular bundle (transverse), μm; 14 – diameter
of vascular bundle (longitudinal), μm;
15 – diameter of xylem vessels (transverse), μm;
16 – diameter of xylem vessels (longitudinal),
μm; 17 – height of phloem, μm; 18 – diameter
of bundle sheath cells, μm; 19 – thickness
of xylem vessels walls, μm; 20 – thickness
of upper epidermis cells outer walls, μm;
21 – thickness of lower epidermis cells outer
walls, μm; 22 – thickness of upper epidermis
cuticle, μm; 23 – thickness of lower epidermis
cuticle, μm; 24 – width of stomata on the
abaxial surface, μm; 25 – width of stomata on
the adaxial surface, μm; 26 – length of stomata
on the abaxial surface, μm; 27 – length of
stomata on the adaxial surface, μm; 28 – leaf
area, dm2; 29 – leaf perimeter, dm; 30 – leaf
mass of defined area, g / dm2; 31 – leaf mass
of a defined volume, g / cm3; 32 – number of
stomata on the abaxial leaf surface, pcs. / mm2;
33 – number of stomata on the adaxial leaf
surface, pcs. / mm2; 34 – number of ordinary
epidermal cells on the abaxial leaf surface,
pcs. / mm2; 35 – number of ordinary epidermal
cells on the adaxial leaf surface, pcs. / mm2;
36 – stomatal index of leaf abaxial surface %;
37 – stomatal index of leaf adaxial surface, %.
Vascular bundles have elliptical form, and
therefore their size and form description
suspect definition of two mutually
perpendicular diameters: longitudinal (the
longest one) and transverse (the shortest one),
which we used for our analysis.
Cluster analysis was performed using
UPGMA method. UPGMA is effective when
objects actually form different “groves”;
however, it works equally well in cases of
extended (chain type) clusters (Systat Software,
2010; Addinsoft, 2017). Simultaneously,
Principle Component Analysis (PCA) was
applied as one of the most frequently used
multivariate data analysis methods suitable for
our purposes (Addinsoft, 2017).
The anatomical structure of the leaves
was investigated using a microscope Nikon
Eclipse E-200 (Japan). Measurements of linear
dimensions and areas were performed using
Image-Pro Premier 9.1 software. Statistica 6.0
and SigmaPlot softwares were used for the
statistical processing of obtained data (Box
et al., 2005; Systat Software, 2010; Everitt et
al., 2011).
Results and discussion
Ecological conditions of the habitat influence
affect morphology and anatomy of plants in
92 Plant Introduction • 89/90
I. Kovalyshyn, A. Pinchuk, A. Likhanov
different ways. In case of studied clematises,
altitude and insolation play principal role. The
common feature of high-altitudinal plants
is relatively thick leaf blades (Kogami et al.,
2001; Zarinkamar et al., 2011; Ahmad et al.,
2016; Arimy et al., 2017). It was also found that
such plants, if compare with the plants from
lower altitudes, have lower stomata density
(Zarinkamar et al., 2011; Arimy et al., 2017),
intensive sclerification around the vascular
bundles (Ahmad et al., 2016), thick walls of
epidermal cells, and, usually, amphistomatic
leaves (Ornellas et al., 2019). From the other
hand, in case of sufficient insolation, neutral
and filtered shades cause the decrease of
leaf thickness and area, leaf weight, stomatal
density, palisade cell length, mesophyll
thickness, and increase in chlorophyll content,
degree of air spaces, adaxial width of palisade
cell, and perimeter to area ratio (Buisson &
Lee, 1993; Hanba et al., 2002).
Leaves of studied clematises were
imparipinnate petiolate, with leaflets
having tripartite (except C. heracleifolia,
which preserves the triple structure
of leaves), toothed (C. alpina ‘Pamela
Jackman’, C. macropetala ‘Maidwell Hall’,
C. ispahanica ‘Zvezdograd’, C fargesii ‘Paul
Farges’, C. tibetana, and C. heracleifolia) or
entire (C. integrifolia ‘Aljonushka’, C. taxensis
‘Princess Diana’, and C. viticella) outlines of
their margins. The leaf tip was always pointed.
The leaf bases varied from wedge-shaped
to notched (Fig. 1). Additionally, climbing
clematises had modified petioles-tendrils for
attachment to the vertical surfaces.
In general, the leaves of the studied plants
differed significantly in shape and area
(Fig. 2). The leaf surface area of the studied
plants ranged from 1838 mm2 in C. integrifolia
‘Aljonushka’ to 18095 mm2 in C. heracleifolia.
The leaf blades of the studied clematises
demonstrated dorsoventral structure. There
were plants with hypostomatic (C. viticella,
C. fargesii ‘Paul Farges’, C. heracleifolia,
C. texensis ‘Princess Diana’, C. macropetala
‘Maidwell Hall’, and C. alpina ‘Pamela Jackman’)
and amphistomatic leaves (C. ispahanica
‘Zvezdograd’ and C. tibetana). In C. integrifolia
‘Aljonushka’, beside the numerous stomata on
the abaxial surface of the leaves, there were
also observed solitary stomata on their adaxial
side. In all studied plants, the stomata of the
anomocytic type were found. The maximal
stomatal area (C. macropetala ‘Maidwell Hall’)
almost twice exceeded the minimal discovered
one (C. texensis ‘Princess Diana’). Ordinary
epidermal cells varied in shape and size and
differed between the experimental plants
and even within the leaf surfaces of the same
taxon. The height of mesophyll and the ratio
of palisade and spongy layers differed among
studied plants too (Fig. 3).
The minimal value of the palisade coefficient
was found in the leaves of C. alpina ‘Pamela
Jackman’ (27.3 %). For the rest plants, it was
within the range of 30–40 % (C. macropetala
‘Maidwell Hall’, C. integrifolia ‘Aljonushka’,
C. fargesii ‘Paul Farges’, C. texensis ‘Princess
Diana’, and C. heracleifolia) or 40–50 %
(C. viticella, C. ispahanica ‘Zvezdograd’, and
C. tibetana).
Palisade mesophyll was represented by two
layers of cells in C. ispahanica ‘Zvezdograd’,
C. integrifolia ‘Aljonushka’, and C. tibetana, and
by a single layer in other studied taxa. Loose
spongy mesophyll with large intercellular
spaces was found in C. texensis ‘Princess
Diana’, C. heracleifolia, C. fargesii ‘Paul Farges’,
and C. tibetana. In other studied plants, spongy
mesophyll was denser with clear boundaries
between the layers. In C. integrifolia
‘Aljonushka’ it consisted of three or four
cell layers, while in C. viticella and C. alpina
‘Pamela Jackman’ – of three layers (Fig. 4).
Leaf morphological and anatomical features
were analysed by PCA. We used nine clematis
taxa as observations and 37 characters and
indexes of the leaf blade as variables (Fig. 5).
From the list of investigated characters, we
selected 22 variables, which made the most
substantial contribution to PCA dispersion.
Selected features in complex allowed
comparing light intensity preferences and
drought-resistance of plants and their ability
to accumulate and retain water (Table 2).
The PCA showed that C. texensis ‘Princess
Diana’ differs from other studied plants by
the thickness of outer cell walls of upper
epidermis, while C. macropetala ‘Maidwell Hall’
differs by the size of cells of spongy mesophyll
and lower epidermis. Clematises with
largest (C. heracleifolia and C. fargesii ‘Paul
Farges’) and smallest leaves (C. integrifolia
‘Aljonushka’), sure thing, strongly differed
from other studied plants by the leaf area.
Besides this, C. integrifolia ‘Aljonushka’ and
C. fargesii ‘Paul Farges’ differed by a high
Plant Introduction • 89/90 93
Morphological features of the leaves in clematises and their ecological peculiarities
BA C
ED F
HG I
Figure 1. Outlines of Clematis leaves: A – C. texensis ‘Princess Diana’; B – C. heracleifolia; C – C. integrifolia
‘Aljonushka’; D – C. fargesii ‘Paul Farges’; E – C. viticella; F – C. alpina ‘Pamela Jackman’; G – C. ispahanica
‘Zvezdograd’; H – C. tibetana; I – C. macropetala ‘Maidwell Hall’.
94 Plant Introduction • 89/90
I. Kovalyshyn, A. Pinchuk, A. Likhanov
Figure 2. The leaf area of studied clematises.
Figure 3. The height of palisade and spongy mesophyll in Clematis leaves.
12231
5514
18095
2406
1838
7978
12052
2157
5326
0 5000 10000 15000 20000
С. texensis 'Princess Diana'
C. macropetala 'Maidwell Hall'
C. heracleifolia
C. ispahanika 'Zvezdograd'
C. integrifolia 'Aljonushka'
C. viticella
C. fargesii 'Paul Farges'
C. alpina 'Pamela Jackman'
C. tibetana
Leaf area, mm2
63.90
74.26
97.48
79.82
63.19
49.11
92.61
66.67
92.53
38.25
40.07
52.70
74.14
36.08
36.31
46.43
25.01
92.11
0 50 100 150 200
C. texensis 'Princess Diana'
C. macropetala 'Maidwell Hall'
C. heracleifolia
C. ispahanica 'Zvezdograd'
C. integrifolia 'Aljonushka'
C. viticella
C. fargesii 'Paul Farges'
C. alpina 'Pamela Jackman'
C. tibetana
Spongy mesophyll Palisade mesophyll
stomata index of the abaxial leaf surface,
and C. heracleifolia – by the low value of this
indicator. Close to C. ispahanica ‘Zvezdograd’,
we traced the accumulation of the features
associated with the stomata apparatus of the
adaxial leaf surface, and thickness of cuticle,
palisade mesophyll, and xylem walls. From
other studied plants, C. tibetana differed in
the thickness of a leaf blade and the height
of lower epidermis cells. PCA projections of
C. viticella and C. alpina ‘Pamela Jackman’
appeared distant from other studied taxa on
the biplot. Following features were found
inherent for C. viticella: the lowest values
of thickness of the leaf blade, diameter of
vascular bundle, diameter of xylem vessels,
and average leaf area. The thinnest palisade
mesophyll and cuticle were found in C. alpina
‘Pamela Jackman’.
The anatomical features of studied
clematises were quite heterogeneous, which
probably, reflects their ecological variability.
In the dendrogram, clematises were
grouped into three main clusters. The first
one included C. tibetana and C. ispahanica
‘Zvezdograd’ with the thickest amphistomatic
leaves. Cultivars C. alpina ‘Pamela Jackman’
and C. macropetala ‘Maidwell Hall’ with
hypostomatic leaves and moderate thickness
of the leaf blade formed the second cluster.
Plant Introduction • 89/90 95
Morphological features of the leaves in clematises and their ecological peculiarities
B
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96 Plant Introduction • 89/90
I. Kovalyshyn, A. Pinchuk, A. Likhanov
Figure 5. PCA biplot of the studied Clematis taxa and their leaf characters. See Table 2 for the character
codes. A – C. texensis ‘Princess Diana’; B – C. macropetala ‘Maidwell Hall’; C – C. heracleifolia; D – C. ispahanica
‘Zvezdograd’; E – C. integrifolia ‘Aljonushka’; F – C. viticella; G – C. fargesii ‘Paul Farges’; H – C. alpina ‘Pamela
Jackman’; I – C. tibetana.
The third and the largest cluster included
C. viticella, C. fargesii ‘Paul Farges’, C.
integrifolia ‘Aljonushka’, C. heracleifolia, and
C. texensis ‘Princess Diana’ with hypostomatic
leaves thickness of which varied from 109.91
to 184.90 μm (Fig. 6).
Despite the difference in origin and
habitus, among studied plants, C. heracleifolia
and C. texensis ‘Princess Diana’ were found
to be the most similar. The leaves of these
plants have the largest area and comparatively
thin epidermis, which indicates their
mesophytic origin. Another subcluster was
formed by the cultivars C. fargesii ‘Paul
Farges’ and C. integrifolia ‘Aljonushka’. Such
species common for Ukraine as C. vitalba
and C. integrifolia were used to create these
cultivars. Hence, these cultivars can occupy
the niche of parental species in the plantations
of deciduous woody plants with low canopy
density. In urban landscapes, they can be used
for growing in containers, to decorate walls
and fences.
The natural range of distribution of
C. viticella is located in southern Europe,
where it grows in the forest ecotons.
Plant Introduction • 89/90 97
Morphological features of the leaves in clematises and their ecological peculiarities
No Variables
Contributions
PC1 PC2 PC3
1 Leaf blade thickness 7.846 0.386 0.759
2 Height of the upper epidermis cells 3.891 0.111 0.026
3 Width of upper epidermis cell 5.616 1.906 1.610
4 Height of lower epidermis cells 7.035 0.037 0.336
5 Width of lower epidermis cells 4.025 5.150 0.027
6 Thickness of palisade mesophyll layer 5.519 2.735 0.379
7 Height of palisade mesophyll cell 4.661 0.139 0.976
8 Width of palisade mesophyll cell 2.848 1.029 2.787
9 Thickness of spongy mesophyll 3.213 0.743 4.589
10 Height of spongy mesophyll cells 1.389 4.546 0.041
11 Width of spongy mesophyll cells 0.149 5.354 0.471
12 Diameter of vascular bundle with sheath cells 0.315 7.050 0.190
13 Diameter of vascular bundle (transverse) 0.358 8.044 1.714
14 Diameter of vascular bundle (longitudinal) 0.940 1.378 6.781
15 Diameter of xylem vessels (transverse) 0.470 2.537 0.407
16 Diameter of xylem vessels (longitudinal) 0.004 2.639 1.643
17 Height of phloem 1.294 1.104 2.538
18 Diameter of bundle sheath cells 1.755 1.007 0.836
19 Thickness of xylem vessel walls 4.170 3.261 1.150
20 Thickness of upper epidermis cells outer walls 4.415 4.416 0.805
21 Thickness of lower epidermis cells outer walls 2.658 5.377 1.125
22 Thickness of upper cuticle 3.008 1.120 2.076
23 Thickness of lower cuticle 2.154 2.663 2.635
24 Width of stomata on the abaxial surface 0.292 1.672 11.160
25 Width of stomata on the adaxial surface 5.508 2.884 0.822
26 Length of stomata on the abaxial surface 5.452 3.678 0.429
27 Length of stomata on the adaxial surface 5.700 2.686 0.421
28 Leaf area 1.016 0.100 10.198
29 Leaf perimeter 0.565 0.265 5.784
30 Leaf mass of defined area 4.116 3.395 0.660
31 Leaf mass of a defined volume 0.418 3.114 4.277
32 Number of stomata on the abaxial leaf surface 0.571 3.079 5.127
33 Number of stomata on the adaxial leaf surface 2.707 3.126 5.809
34 Number of ordinary epidermal cells on the abaxial leaf surface 0.935 3.531 10.122
35 Number of ordinary epidermal cells on the adaxial leaf surface 0.052 5.142 8.596
36 Stomatal index of leaf abaxial surface 0.129 1.398 0.543
37 Stomatal index of leaf adaxial surface 4.806 3.198 2.153
Table 2. Contribution of selected characters to PCA dispersion. The characters making a significant
contribution are indicated in bold.
98 Plant Introduction • 89/90
I. Kovalyshyn, A. Pinchuk, A. Likhanov
Figure 6. UPGMA clustering of studied clematises basing on morpho-anatomical parameters.
This explains the shade tolerance peculiarities
in the structure of its leaves. Therefore, this
species, as a bright summer accent, fits the
woody plantations with low canopy density.
Cultivars C. alpina ‘Pamela Jackman’
and C. macropetala ‘Maidwell Hall’ are
traditionally attributed to the Atragene
group, which combines clematises from
the Northern Hemisphere forests. These
plants grow in partial shade, and bloom
in spring during rapid fluctuations of the
daily temperature under a sparse canopy of
the trees in windy forest biocenoses. Small
leaves and their dissection can be a result
of adaptation of these plants to mentioned
growing conditions. Hence, the best places
for their growth in the city are on the edges
and along the paths in the parks and squares
composed by deciduous trees. In such
conditions they will get enough light during
the flowering period and enough shade
during the hot summer time.
Stony steppe slopes with high insolation
are the natural habitats of C. tibetana
and C. ispahanica. Clematis tibetana and
C. ispahanica Boiss. ‘Zvezdograd’ have
the following leaf structure features:
amphistomatic leaves, high palisade
coefficient, and relatively significant total
leaf blade thickness. Such features are crucial
for the plants growing on mountain slopes
with high insolation and fluctuations in
daily temperatures. The small area of leaves
allows these plants to avoid damage by
ground particles during the winds. While the
relatively thick cuticle reduces transpiration
and refracts sunlight. These traits are
genetically determined and persist even in
plants growing in urban areas. Therefore,
C. tibetana and C. ispahanica ‘Zvezdograd’
are appropriate for growing on windy, rocky
mounds.
Conclusions
Adaptive features of the studied clematises
are different. Complex analysis of their
leaf morpho-anatomical structure allowed
assessing the ecological preferences of
these plants. In particular, among the main
characters of heliophilous clematises were
determined amphistomatic leaves, a high
palisade coefficient, and a considerably
thick cuticle. The small diameter of vascular
bundles and the thin cuticle layer are signs
of shade-tolerant clematises. While the
mesophytic clematises combine a thin cuticle
layer and large size of stomata, mesophilic
and epidermal cells. Small-celled leaf tissues,
in general, indicate drought-resistant
clematises.
5 6 7 8 9 10 11 12 13
C. tibetana
C. ispahanica `Zvezdograd`
C. alpina `Pamela Jackman`
C. macropetala `Maidwell Hall`
C. viticella
C. fargesii `Paul Farges`
C. integrifolia `Aljonushka`
C. heracleifolia
C. texensis `Princess Diana`
Plant Introduction • 89/90 99
Morphological features of the leaves in clematises and their ecological peculiarities
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100 Plant Introduction • 89/90
I. Kovalyshyn, A. Pinchuk, A. Likhanov
Морфологічні ознаки листків ломиносів як відображення їх екологічних
особливостей
Ірина Ковалишин 1, *, Андрій Пінчук 2, Артур Ліханов 2
1 Інститут фізіології рослин і генетики Національної академії наук України, вул. Васильківська, 31/17,
Київ, 03022, Україна; * Iryna_B_Kovalyshyn@ukr.net
2 Національний університет біоресурсів і природокористування України, вул. Героїв Оборони, 15,
Київ, 03041, Україна
Визначення кількісних показників морфологічних і анатомічних ознак листкових пластинок
здійснювали для дев’яти видів і сортів ломиносів (Clematis alpina ‘Pamela Jackman’, C. macropetala
‘Maidwell Hall’, C. integrifolia ‘Aljonushka’, C. ispahanica ‘Zvezdograd’, C. fargesii ‘Paul Farges’, C. texensis
‘Princess Diana’, C. tibetana, C. viticella та C. heracleifolia).
Серед досліджених ломиносів були присутні рослини з гіпостоматичними (C. viticella, C. fargesii ‘Paul
Farges’, C. heracleifolia, С. texensis ‘Princess Diana’, C. macropetala ‘Maidwell Hall’, C. alpina ‘Pamela Jackman’) та
амфістоматичними листками (C. ispahanica ‘Zvezdograd’ і C. tibetana). А також рослини з гіпостоматичні
листками, які мали поодинокі продихи на адаксіальній поверхні листків (C. integrifolia ‘Aljonushka’).
Коефіцієнт палісадності листків коливався від 27,3 % (C. alpina ‘Pamela Jackman’) до 49,9 % (C. tibetana).
Листки досліджених рослин також суттєво відрізнялися за площею, зокрема у C. integrifolia ‘Aljonushka’
площа листкової поверхні майже у десять разів перевищувала таку у C. heracleifolia.
У результаті кластерного аналізу, перший кластер був сформований C. tibetana та C. ispahanica
‘Zvezdograd’. Ці рослини, здатні виживати в умовах різкого перепаду температур і вітряної погоди
на відкритих кам’янистих ділянках. До другого кластеру увійшли C. alpina ‘Pamela Jackman’ та
C. macropetala ‘Maidwell Hall’ – рослини, які цвітуть навесні, в період значної зміни добових температур.
Порівняно невелика площа листків цих рослин може вказувати на адаптацію шляхом зменшення
транспіраційної площі та парусності. Третій кластер об’єднав мезофітні рослини, які відрізняються
від решти більшою загальною площею листків. Однак, до третього кластеру також увійшов
культивар C. integrifolia ‘Aljonushka’, який відрізняється найменшою площею листкової пластинки,
але має подібну морфо-анатомічну будову.
Ключові слова: Clematis, кластерний аналіз, листок, епідерма, мезофіл
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|
| id | oai:ojs2.plantintroduction.org:article-1564 |
| institution | Plant Introduction |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2025-07-17T12:53:42Z |
| publishDate | 2021 |
| publisher | M.M. Gryshko National Botanical Garden of the NAS of Ukraine |
| record_format | ojs |
| resource_txt_mv | wwwplantintroductionorg/e1/2c00b9ab0aa818442eae798f746833e1.pdf |
| spelling | oai:ojs2.plantintroduction.org:article-15642023-08-26T20:39:20Z Morphological features of the leaves in clematises as a reflection of their ecological peculiarities Морфологічні ознаки листків ломиносів як відображення їх екологічних особливостей Kovalyshyn, Iryna Pinchuk, Andrii Likhanov, Artur Quantitative morpho-anatomical features of leaves of nine Clematis taxa (C. alpina ‘Pamela Jackman’, C. macropetala ‘Maidwell Hall’, C. integrifolia ‘Aljonushka’, C. ispahanica ‘Zvezdograd’, C. fargesii ‘Paul Farges’, C. texensis ‘Princess Diana’, C. tibetana, C. viticella, and C. heracleifolia) were determined with the aim to analyze their adaptation to the environmental conditions.Among investigated clematises, there were plants with hypostomatic (C. viticella, C. fargesii ‘Paul Farges’, C. heracleifolia, C. texensis ‘Princess Diana’, C. macropetala ‘Maidwell Hall’, and C. alpina ‘Pamela Jackman’), and amphistomatic leaves (C. ispahanica ‘Zvezdograd’ and C. tibetana). In C. integrifolia ‘Aljonushka’ leaves were hypostomatic, but few solitary stomata were also present on the adaxial surface. In the leaves of investigated taxa, the palisade coefficient ranged from 27.3% (C. alpina ‘Pamela Jackman’) to 49.9% (C. tibetana). The leaves also differed significantly in size. In particular, leaves of C. integrifolia ‘Aljonushka’ were almost ten times smaller than such of C. heracleifolia.As a result of UPGMA clustering, the plants that can survive in severe windy weather in open rocky areas, Clematis tibetana and C. ispahanica ‘Zvezdograd’, were joined in a separate cluster. The second cluster combined C. alpina ‘Pamela Jackman’ and C. macropetala ‘Maidwell Hall’ – cultivars blooming in the spring, during a period of significant difference in daily temperatures. A relatively small leaf area in plants from these two clusters may indicate an adaptation by reducing the transpiration area and general windage. The third cluster united the rest of investigated taxa, mostly – the mesophytic plants with a relatively large leaf area. However, due to similar morpho-anatomical structure of the leaf, the third cluster also comprised C. integrifolia ‘Aljonushka’ with the smallest leaves. Визначення кількісних показників морфологічних і анатомічних ознак листкових пластинок здійснювали для дев’яти видів і сортів ломиносів (Clematis alpina ‘Pamela Jackman’, C. macropetala ‘Maidwell Hall’, C. integrifolia ‘Aljonushka’, C. ispahanica ‘Zvezdograd’, C. fargesii ‘Paul Farges’, C. texensis ‘Princess Diana’, C. tibetana, C. viticella та C. heracleifolia).Серед досліджених ломиносів були присутні рослини з гіпостоматичними (C. viticella, C. fargesii ‘Paul Farges’, C. heracleifolia, С. texensis ‘Princess Diana’, C. macropetala ‘Maidwell Hall’, C. alpina ‘Pamela Jackman’) та амфістоматичними листками (C. ispahanica ‘Zvezdograd’ і C. tibetana). А також рослини з гіпостоматичні листками, які мали поодинокі продихи на адаксіальній поверхні листків (C. integrifolia ‘Aljonushka’). Коефіцієнт палісадності листків коливався від 27,3 % (C. alpina ‘Pamela Jackman’) до 49,9 % (C. tibetana). Листки досліджених рослин також суттєво відрізнялися за площею, зокрема у C. integrifolia ‘Aljonushka’ площа листкової поверхні майже у десять разів перевищувала таку у C. heracleifolia.У результаті кластерного аналізу, перший кластер був сформований C. tibetana та C. ispahanica ‘Zvezdograd’. Ці рослини, здатні виживати в умовах різкого перепаду температур і вітряної погоди на відкритих кам’янистих ділянках. До другого кластеру увійшли C. alpina ‘Pamela Jackman’ та C. macropetala ‘Maidwell Hall’ – рослини, які цвітуть навесні, в період значної зміни добових температур. Порівняно невелика площа листків цих рослин може вказувати на адаптацію шляхом зменшення транспіраційної площі та парусності. Третій кластер об’єднав мезофітні рослини, які відрізняються від решти більшою загальною площею листків. Однак, до третього кластеру також увійшов культивар C. integrifolia ‘Aljonushka’, який відрізняється найменшою площею листкової пластинки, але має подібну морфо-анатомічну будову. M.M. Gryshko National Botanical Garden of the NAS of Ukraine 2021-06-14 Article Article application/pdf https://www.plantintroduction.org/index.php/pi/article/view/1564 10.46341/PI2021002 Plant Introduction; No 89/90 (2021); 89-100 Інтродукція Рослин; № 89/90 (2021); 89-100 2663-290X 1605-6574 10.46341/PI89-90 en https://www.plantintroduction.org/index.php/pi/article/view/1564/1513 Copyright (c) 2021 Iryna Kovalyshyn, Andrii Pinchuk, Artur Likhanov http://creativecommons.org/licenses/by/4.0 |
| spellingShingle | Kovalyshyn, Iryna Pinchuk, Andrii Likhanov, Artur Морфологічні ознаки листків ломиносів як відображення їх екологічних особливостей |
| title | Морфологічні ознаки листків ломиносів як відображення їх екологічних особливостей |
| title_alt | Morphological features of the leaves in clematises as a reflection of their ecological peculiarities |
| title_full | Морфологічні ознаки листків ломиносів як відображення їх екологічних особливостей |
| title_fullStr | Морфологічні ознаки листків ломиносів як відображення їх екологічних особливостей |
| title_full_unstemmed | Морфологічні ознаки листків ломиносів як відображення їх екологічних особливостей |
| title_short | Морфологічні ознаки листків ломиносів як відображення їх екологічних особливостей |
| title_sort | морфологічні ознаки листків ломиносів як відображення їх екологічних особливостей |
| url | https://www.plantintroduction.org/index.php/pi/article/view/1564 |
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