Карпологічні особливості деяких представників роду Artemisіa, поширених в Україні
The achenes’ micromorphological structure of six Artemisia species (A. absinthium, A. annua, A. argyi, A. dracunculus, A. marschalliana, and A. vulgaris) distributed in Ukraine was investigated with the aim to identify taxonomically significant carpological features for the studied species.Fruits we...
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M.M. Gryshko National Botanical Garden of the NAS of Ukraine
2020
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Plant Introduction| _version_ | 1860145096062664704 |
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| author | Vakulenko, Tetyana Bagatska, Tetyana Korablova, Olga Gurnenko, Ivan Rakhmetov, Dzhamal |
| author_facet | Vakulenko, Tetyana Bagatska, Tetyana Korablova, Olga Gurnenko, Ivan Rakhmetov, Dzhamal |
| author_sort | Vakulenko, Tetyana |
| baseUrl_str | https://www.plantintroduction.org/index.php/pi/oai |
| collection | OJS |
| datestamp_date | 2023-08-26T20:39:33Z |
| description | The achenes’ micromorphological structure of six Artemisia species (A. absinthium, A. annua, A. argyi, A. dracunculus, A. marschalliana, and A. vulgaris) distributed in Ukraine was investigated with the aim to identify taxonomically significant carpological features for the studied species.Fruits were collected from wild plants growing in the recreation areas of Kyiv and from the plants hosted in the living collection of the M.M. Gryshko National Botanical Garden of the NAS of Ukraine. Macromorphological features were studied with a binocular microscope Stemi 2000 Carl Zeiss. The ultrastructure of the surface was studied using a scanning electron microscope JSM-6700F JEOL, in the mode of secondary electron emission. Carbon spraying was performed using vacuum thermal evaporation of the vacuum universal post VUP-5M SELMI; platinum spraying was performed with an ion-plasma etching device JFC-1600 JEOL.All investigated species have a well-expressed, true, multicellular, symmetrical, multi-row carpopodium but differ by its shape. The surface of A. annua achenes is reticulate-pitted due to the specific structure of the exocarp and the exotests located below it. Species with lobed and continuous apical rim and species without a rim around the place of attachment of the corolla were found. It was established that the primary ultrastructure of investigated fruits is caused by the shape and character of mucous cell arrangement; epidermal cells of the exocarp are mostly compressed, without clear boundaries. Mostly a thin and inconspicuous cuticle, sometimes wrinkled, with layers and small outgrowths, causes the secondary ultrastructure of these fruits. We believe that the mentioned carpological features may be used as additional criteria in combination with other features to identify studied species. |
| doi_str_mv | 10.46341/PI2020038 |
| first_indexed | 2025-07-17T12:53:47Z |
| format | Article |
| fulltext |
© The Authors. This content is provided under CC BY 4.0 license.
Plant Introduction, 87/88, 54–64 (2020)
RESEARCH ARTICLE
Carpological features of certain Artemisia species distributed in Ukraine
Introduction
The genus Artemisia L. is the largest in
the tribe Anthemideae Cass. of the family
Asteraceae Dumort., reaching up to 550
species spread throughout the Northern
Hemisphere (Mungalov, 2004; Beer, 2005;
Boyko, 2011). There are 30 Artemisia species
distributed in Ukraine (Boyko, 2013). The
systematics of this genus is complicated
due to significant intraspecific variability
of morphological features. Such a situation
encourages searching for new diagnostic
criteria. Carpological features may be among
the solutions because their importance for the
taxonomy and phylogeny of Artemisia has been
proved (Boyko, 2011; Boyko & Novozhilova,
2018; Bagatskaya & Vakulenko, 2013).
Artemisia fruit is an indehiscent achene
that develops from the inferior bicarpellate
(dimerous) ovary. The mature seed cover
is formed by three structural elements:
the pericarp, the spermoderm, and the
endosperm’s remnants. The pericarp in
Tetyana Vakulenko *, Tetyana Bagatska, Olga Korablova **, Ivan Gurnenko,
Dzhamal Rakhmetov
M.M. Gryshko National Botanical Garden, National Academy of Sciences of Ukraine, Tymiryazyevska str. 1, 01014 Kyiv, Ukraine;
* botanicukr@gmail.com, ** okorablova@ukr.net
Received: 28.10.2020 | Accepted: 21.12.2020 | Published: 30.12.2020
Abstract
The achenes’ micromorphological structure of six Artemisia species (A. absinthium, A. annua, A. argyi,
A. dracunculus, A. marschalliana, and A. vulgaris) distributed in Ukraine was investigated with the aim to
identify taxonomically significant carpological features for the studied species.
Fruits were collected from wild plants growing in the recreation areas of Kyiv and from the plants hosted in the
living collection of the M.M. Gryshko National Botanical Garden of the NAS of Ukraine. Macromorphological
features were studied with a binocular microscope Stemi 2000 Carl Zeiss. The ultrastructure of the surface
was studied using a scanning electron microscope JSM-6700F JEOL, in the mode of secondary electron
emission. Carbon spraying was performed using vacuum thermal evaporation of the vacuum universal
post VUP-5M SELMI; platinum spraying was performed with an ion-plasma etching device JFC-1600 JEOL.
All investigated species have a well-expressed, true, multicellular, symmetrical, multi-row carpopodium
but differ by its shape. The surface of A. annua achenes is reticulate-pitted due to the specific structure
of the exocarp and the exotests located below it. Species with lobed and continuous apical rim and
species without a rim around the place of attachment of the corolla were found. It was established that
the primary ultrastructure of investigated fruits is caused by the shape and character of mucous cell
arrangement; epidermal cells of the exocarp are mostly compressed, without clear boundaries. Mostly
a thin and inconspicuous cuticle, sometimes wrinkled, with layers and small outgrowths, causes the
secondary ultrastructure of these fruits. We believe that the mentioned carpological features may be used
as additional criteria in combination with other features to identify studied species.
Keywords: Asteraceae, Artemisia, carpopodium, mucous complexes, primary ultrastructure, secondary ultrastructure, fruit
https://doi.org/10.46341/PI2020038
UDC 581.47+581.16
https://creativecommons.org/licenses/by/4.0/
https://orcid.org/0000-0002-6141-6689
https://orcid.org/0000-0001-6661-939X
https://orcid.org/0000-0001-7600-6909
https://orcid.org/0000-0001-6656-4640
https://orcid.org/0000-0001-7260-3263
Plant Introduction • 87/88 55
Carpological features of certain Artemisia species distributed in Ukraine
Artemisia is quite simplified, formed by a
single layer of epidermal cells, remnants of
vascular bundles, and an additional layer of
subepidermal cells (Boyko, 2011).
The exocarp of Artemisia fruit contains
mucous cells. Mucous cells are located solitary
or in the so-called ‘mucous complexes’ forming
longitudinal raised sleeper-like or ladder-
shaped ribs with tangentially elongated cells
(Yakovleva et al., 2002). The cells of mucous
complexes differ significantly from the regular
epidermal cells. Such complexes can be of
different length, sometimes even as long as
achene (Boyko, 2011).
Scanning electron microscopy (SEM)
is extremely important for the fruit
micromorphology. It allows to study the
ultrastructure of integuments in more details
and to elucidate additional carpological
features such as the structure of the
carpopodium (the area of separation of the
achene from the perianth placed around the
fruit scar), the structure of the apical bowl (the
area of attachment of the corolla, surrounded
by a roller or rim), and primary and secondary
ultrastructure of the surface (Mukherjee
& Nordenstam, 2010; Boyko, 2011). SEM of
the fruits often helps to resolve taxonomic
issues. Hence, the work aimed to study the
micromorphology of achenes of six Artemisia
species represented in the flora of Ukraine and
determine carpological features that could be
applied to distinguish these taxa.
Material and methods
Mature achenes of Artemisia absinthium L.,
A. annua L., A. argyi Lévl. et Vaniot,
A. dracunculus L., A. marschalliana Spreng,
and A. vulgaris L. were studied. Achenes
were collected from the wild plants growing
in the recreation areas of Kyiv (two localities
– 50.511839 N, 30.399793 E and 50.514308 N,
30.387294 E), as well as from the collection of
aromatic plants at the M.M. Gryshko National
Botanical Garden of the NAS of Ukraine
(50.416432 N, 30.557891 E). All the investigated
localities are in the same Forest-Steppe zone
of Ukraine.
Macromorphological features were
analyzed under light microscopy (LM) using a
binocular microscope Stemi 2000, Carl Zeiss
(Germany). Digital processing of images was
carried out in the program Axio Vision. The
ultrastructure of the surface was studied using
a scanning electron microscope JSM-6700F,
JEOL (Japan) in the mode of secondary electron
emission. Carbon spraying was performed by
the method of vacuum thermal evaporation
of the vacuum universal post VUP-5M, SELMI
(Ukraine). Platinum spraying was performed
using an ion-plasma etching device, JFC-1600,
JEOL (Japan).
General fruit descriptions are provided
following Artyushenko & Fedorov (1986) and
Bojnanský & Fargašová (2007). To characterize
the ultrastructure, we used Barthlott (1981)
terminology, according to which the difference
between primary and secondary structure
can be observed. Applied terminology has
also been crosschecked accordingly to Stern
(1992). Consequently, it was accepted that
macromorphological features (shape of
epidermal cells, curvature, height, thickness
and relief of their periclinal and anticlinal walls,
the origin of indumentum, etc.) determine
the primary structure of the fruits in studied
species. By contrast, the secondary structure
is determined by the micromorphology and
characterizes the cuticle sculpture.
Results
Artemisia absinthium
Achenes are oblong-wedge-shaped, slightly
curved dorsally, 0.93–1.07 × 0.37–0.45 mm,
planar compressed-polygonal. Ribs smoothed,
weakly defined. The surface is bare, glossy;
the relief is longitudinally striped, the color
is brown or light brown (Fig. 1 I). The tip
is widened, slightly beveled, the place of
attachment of the corolla with a diameter of
220–250 μm, surrounded by a low blade rim
(Fig. 2 I – B). The base is narrow, carpopodium
basal, whitish, triangular-rounded, 80 ×
110 μm, not wrapped, with four-five rows
of cells. Carpopodial cells 15–25 × 5–12 μm,
tangentially elongated, rectangular in the
lower rows, often polygonal in the upper
rows, due to which the dividing line between
the carpopodium and the achene body is
wavy (Fig. 2 I – A). The primary ultrastructure
of the carpopodium is well defined, the cell
boundaries are clearly visible, and anticline
walls are thin, straight, or slightly rounded.
The outer periclinal walls are convex.
56 Plant Introduction • 87/88
T. Vakulenko, T. Bagatska, O. Korablova, I. Gurnenko, D. Rakhmetov
Figure 1. General view of Artemisia fruits under the light (LM) and scanning electron (SEM) microscopes:
I – A. absinthium; II – A. annua; III – A. argyi; IV– A. dracunculus; V – A. marschalliana; VI – A. vulgaris.
LM SEM
I
II
III
IV
V
VI
Plant Introduction • 87/88 57
Carpological features of certain Artemisia species distributed in Ukraine
A B
I
II
III
IV
V
VI
Figure 2. SEM structure of carpopodium (A) and apical bowl (B) of Artemisia species: I – A. absinthium;
II – A. annua; III – A. argyi; IV– A. dracunculus; V – A. marschalliana; VI – A. vulgaris.
58 Plant Introduction • 87/88
T. Vakulenko, T. Bagatska, O. Korablova, I. Gurnenko, D. Rakhmetov
A smooth cuticle represents the secondary
ultrastructure.
Mucous complexes in the form of
longitudinal single-row strands consist of
mucous cells. Most of the cords rise above the
achene’s surface, extending from the apex to
the base, but there are also short complexes
containing 15–30 mucous cells (Fig. 3). The
primary ultrastructure of the strands is
well-visible – they consist of rectangular
tangentially elongated cells up to 15–25 μm
long, tightly spaced one below the other
(Fig. 4 A). Anticline walls of mucous cells
are straight, thickened, with raised edges.
B
D
F
A
C
E
Figure 3. Fragments of the surface of achenes of species of the genus Artemisia with complexes of mucous
cells under the SEM: I – A. absinthium; II – A. annua; III – A. argyi; IV– A. dracunculus; V – A. marschalliana;
VI – A. vulgaris.
Plant Introduction • 87/88 59
Carpological features of certain Artemisia species distributed in Ukraine
Outer periclinal walls – concave or almost
flat, densely covered with longitudinal folds.
The exocarp’s epidermal cells, which do not
contain mucus, situate between the strands
of mucous complexes in two-three rows.
According to the primary ultrastructure, they
are longitudinally elongated, prosenchymatic,
with almost straight or tortuous thickened
raised anticline walls and concave outer
periclinal, often folded or grooved. In some
places, the primary ultrastructure is not well-
defined, and the cell boundaries are weakly
visible. The secondary ultrastructure is weakly
expressed – the cuticle is mostly smooth,
occasionally slightly wavy, with isolated bumpy
cuticular outgrowths.
Artemisia annua
Achenes are pear-shaped, 0.75–0.85 × 0.39–
0.45 mm, slightly curved. The surface is bare,
glossy; the relief is fine-mesh, with narrow
longitudinal ribs, straw-brown color, straw-
yellowish (Fig. 1 II). The top is widened and
rounded. The place of attachment of the
corolla is slightly bent on the ventral side,
rounded, 100–120 μm in diameter, often with
remnants of stylopodium in the center in the
form of a whitish column, surrounded by a
low solid rim (Fig. 2 B – II). The base is narrow,
straight, or slightly beveled. Carpopodium is
basal, round, annular, symmetrical, up to 50–
75 μm in diameter, not wrapped. Carpopodial
cells are tangentially elongated, 10–15 ×
4-8 μm, upper row cells are occasionally
polygonal. The demarcation line between the
carpopodium and the achene body is slightly
wavy (Fig. 2 A – II). The primary ultrastructure is
well-defined; the anticline walls of carpopodial
cells are thin and straight; external periclinal
walls – convex and thickened. A smooth cuticle
represents the secondary ultrastructure.
A B
Figure 4. The strands of the mucous cells of Artemisia
absinthium (A) and A. marschalliana (B) under the LM.
Mucous cells are grouped in complexes
that extend in single-row strands along
the achene, barely rising above its body.
The strands of mucous complexes can be
short, sometimes bifurcated, or, conversely,
combined into one. These strands have wavy
contours formed by tangentially elongated
mucous cells of different lengths (from 20 to
40 μm). The primary ultrastructure of the
strands is relatively well-defined. Anticlinal
walls are significantly raised, mostly wavy
and indistinct. External periclinal walls are
concave or almost flat, permeated with
branched more or less rounded folds, forming
a reticulate-pitted relief (Fig. 3). Exocarp cells
are rectangular or prismatic, compressed by
mucous strands. The primary ultrastructure is
poorly visible, anticline walls with significantly
raised wavy edges. External periclinal walls are
often concave, sink into the cells’ cavities; their
folds are weakly noticeable. The secondary
ultrastructure is defined differently in various
parts of the achene. It is longitudinally wrinkled
near the place of attachment of the corolla
and around the carpopodium. In some places,
there are cuticular bumps, in other places –
often without noticeable differentiation.
Artemisia argyi
Achenes are fusiform, with a straight
truncated-narrowed apex 1.34–1.45 × 0.39–
0.50 mm, rounded-depressed in cross-
section. The surface is longitudinally
wrinkled, bare, and glossy, with several
light ribs, dark brown (Fig. 1 III). The place
of attachment of the corolla is rounded or
compressed-rounded, with a diameter of
up to 210–250 μm, in the center with the
remnants of stylopodium, surrounded by
a low solid rim (Fig. 2 B – III). The base is
narrow, sometimes obliquely cut or turned to
the side. Carpopodium basal, light yellowish,
up to 120–150 μm in diameter, slightly wider
than the base, symmetrical, round, wrapped;
consists of five-seven rows of cells. The line
of demarcation between the carpopodium
and the achene body is clear, almost smooth;
the transition zone between the carpopodium
and the rest of the achene is often twisted
(Fig. 2 A – III). The primary ultrastructure
is well-defined, carpopodial cells are
tangentially elongated, 15–25 × 8–10 μm,
with clear boundaries. Anticlinal walls are
60 Plant Introduction • 87/88
T. Vakulenko, T. Bagatska, O. Korablova, I. Gurnenko, D. Rakhmetov
thin, straight, or slightly curved. The outer
periclinal walls are convex, thickened. The
secondary ultrastructure is not pronounced;
the cuticular relief is smooth.
Mucous complexes form short, sometimes
two-row longitudinal strands, none of which
extends from the base to the top. Mucous
cells are tangentially elongated to 10–15 μm
and are barely raised over the strands’ surface
(Fig. 3). Their primary ultrastructure is well
visible only in a few places. The anticline
walls are thickened, straight. The outer
periclinal walls are flat or concave, often
with a rounded tubercle in the center. The
exocarp’s epidermal cells are 4–5-angular,
longitudinally elongated, significantly
compressed, and often smooth. Their borders
are the best manifested on the ventral side;
anticlinal walls are thickened, with raised
edges, periclinal – concave, rarely flat,
sometimes chaotically folded. Ribs formed by
the vascular bundles complicate the primary
ultrastructure. Schizogenic secretory
containers are present. The secondary
ultrastructure is poorly visible, only in a few
places. The cuticular layer is slightly wavy,
with occasionally noticeable cuticular bumps.
Artemisia dracunculus
Achenes are pear-shaped, 1.37–1.52 × 0.65–
0.77 mm, elliptical in cross-section. The top is
bluntly rounded, sometimes slightly beveled
(Fig. 1 IV). The fastening of a corolla is not
outlined, inclined towards the ventral party
(Fig. 2 B – IV). The ribs are indistinct. In some
places, they are entirely smooth, brownish,
and reddish-brown; the ventral rib is the
most well-defined. The surface is shiny, bare;
the relief is longitudinally wrinkled; the color
is dark brown. Carpopodium basal, straight
or slightly curved on the ventral side, 150 ×
200 μm, oval, symmetrical, wrapped, with
five-six rows of cells. Cells are tangentially
elongated, 20–25 × 10–15 μm, quadrangular,
occasionally polygonal in the upper row. The
demarcation line is clear, almost equal along all
its length (Fig. 2 A – IV). The primary structure
is less pronounced than in previous species.
The boundaries of carpopodial cells are
seen only partly in the upper rows, while the
lower rows remain smoothed cell contours.
Anticlinal walls are thin, straight, or slightly
curved, the outer periclinal – thickened,
convex. The secondary ultrastructure of the
cuticle is straightened, smooth.
Mucous complexes represent smooth,
different length longitudinal strands that are
flush with the achene body and undefined
in some places (Fig. 3). Mucous cells are
tangentially elongated; anticline walls are
slightly thickened, straight. The outer
periclinal walls are concave or almost flat,
sometimes chaotically folded, especially
around the apex. Chaotic cuticular layers
represent the secondary ultrastructure. The
primary structure of the epidermal cells of the
exocarp is also weakly defined. Only in some
places, solitary prosenchymatic cells with
thickened curved anticlinal walls and concave
periclinal are visible. The ribs are more or less
visible in the upper part and at the base of
the achenes; occasionally, above them, there
are located schizogenic secretory containers.
The secondary ultrastructure is indistinct; the
wrinkled surface of the cuticle appears only in
some places.
Artemisia marschalliana
Achenes are inverted ovate, rounded in cross-
section, 0.80–1.05 × 0.37–0.47 mm, sometimes
slightly curved in the dorsal-ventral
projection, with more convex backside, and
the ventral part that is concave. The surface
is bare, slightly shiny, the ribs are smooth,
and the relief is longitudinally striped, with
dark brown color (Fig. 1 V). The apex is
wide, bluntly rounded, often obliquely cut.
The corolla’s attachment place is rounded,
with a diameter of 80–90 μm, surrounded
by a low light rim (Fig. 2 B – V). The base is
narrow. Basal carpopodium, slightly bent to
the side, whitish, annular, symmetrical, and
wrapped, consists of six-seven rows of cells.
Carpopodial cells are tangentially elongated,
rectangular, of different lengths, 13–25 ×
7–12 μm. The upper row often consists of
polygonal cells, due to which the line of
demarcation between the carpopodium and
the achene body is wavy (Fig. 2 A – V). At the
primary ultrastructure level, the anticline
walls of carpopodial cells are straight or
curved and thin, while the outer periclinal
walls are thickened and convex. The cuticle is
without noticeable differentiation.
Mucous complexes are in the form of
longitudinal, slightly raised strands extending
Plant Introduction • 87/88 61
Carpological features of certain Artemisia species distributed in Ukraine
from the base to the top. Mucous cells are
rectangular, 10–25 μm long, placed by two-
three side by side, so the strands are wider
than in other species (Fig. 3). Their anticlinal
walls are straight, slightly thickened;
the outer periclinal – concave in the cell
cavity or smooth, sometimes in the middle
with a tubercle (Fig. 4 B). The secondary
ultrastructure of mucous cells is often
characterized by a straightened cuticle, which
is slightly transversely or longitudinally-
wrinkled around the apex and the base.
The exocarp’s epidermal cell boundaries are
poorly visible, preferably from the achene’s
ventral side, where they are elongated along
the central axis. Anticline walls are more or
less straight, with raised fine-toothed edges.
The outer periclinal wall of epidermal cells is
concave, with many chaotic folds, which often
overlap, hiding the primary ultrastructure.
The ribs are smooth; occasionally, schizogenic
secretory containers are located on them.
The secondary microstructure is not clearly
defined; flat or wrinkled plates represent
cuticular layers.
Artemisia vulgaris
Achenes are fusiform, 1.50–1.60 × 0.40–
0.47 mm, oval in section, with a narrowed
tip and base. The surface is glossy, bare,
longitudinally wrinkled, brown or dark
brown, with light yellowish ribs (Fig. 1 VI).
The place of attachment of the corolla is well
visible, situated slightly to the side, round,
surrounded by a high thin solid translucent
rim up to 150 μm in diameter (Fig. 2 B – VI).
Basal carpopodium, placed directly or slightly
moved to the side, oval, symmetrical, wrapped,
90–100 × 140–150 μm, and consists of seven-
eight rows of cells. Carpopodial cells are
tangentially elongated, 10–12 × 4–8 μm; the
upper row is often made of polygonal cells;
the demarcation line is clearly visible, wavy
(Fig. 2 A – VI). The primary ultrastructure
of the carpopodium in the lower rows is
smooth, without clear boundaries. It is better
defined in the upper rows – the anticlinal
walls are straight, thin, the outer periclinal are
thickened, slightly convex, sometimes flat. The
secondary ultrastructure is not defined; the
cuticle is straightened.
Mucous cells are joined in small separate
groups or complexes in the form of short
cords, located almost on the same level as
the achenes’ body (Fig. 3). Mucous cells in
the strands are sometimes placed by two
in a row. Their primary ultrastructure is
indistinct. Thickened straight anticlinal walls
have raised edges. Concave outer periclinal
walls are sometimes chaotically folded. The
exocarp epidermis cells are located between
the mucous complexes in three-four or
more rows, with thickened, mostly straight
anticlinal walls. The outer periclinal walls are
concave or flat, sometimes convex, sometimes
folded. Above the vascular bundles, there are
sometimes schizogenic secretory containers.
The secondary ultrastructure of exocarp cells
is weakly defined; only in certain places, mainly
at the ends, the transversely wrinkled cuticle
is seen, with occasional cuticular tubercles.
Discussion
Our studies showed that the achenes of the
described species have structural elements
that are typical to other Asteraceae members
(Boyko, 2011). Three types of achenes were
initially distinguished for the genus Artemisia:
narrow-lanceolate, narrowly elliptical, and
inverted ovate (Korobkov, 1973), however
later, the list was expanded (Ouyahaya,
1995; Beer, 2005; Boyko, 2011). Inverted
ovate (A. marschalliana), pear-shaped
(A. dracunculus and A. annua), spindle-shaped
(A. argyi and A. vulgaris), and narrow-wedge-
shaped (A. absinthium) achenes were found.
All studied species have a well-developed
carpopodium consisting of thick-walled cells,
which differ from the regular cells of the
achene. Carpopodium is true, multicellular,
annular, symmetrical, multiline, with four–
eight rows of tangentially elongated cells
with thickened convex outer periclinal walls
and more or less straight anticlinal ones.
Previously, we described a false carpopodium
with concave outer periclinal walls for A. argyi
and A. vulgaris (Bagatskaya & Vakulenko, 2013).
More advanced SEM technique has revealed
in these species a true carpopodium with
convex outer periclinal walls. This discrepancy
may be related to the achenes’ location in the
calathium, the degree of their maturity and
quality, as well as insufficient resolution of the
light microscope (Boyko, 2011). Carpopodium in
the studied species is round (A. annua, A. argyi,
62 Plant Introduction • 87/88
T. Vakulenko, T. Bagatska, O. Korablova, I. Gurnenko, D. Rakhmetov
and A. marschalliana), oval (A. dracunculus
and A. vulgaris), or triangular-round
(A absinthium). In A. annua, A. absinthium,
and A. vulgaris, carpopodial cells are located
only on the outer surface of the achene, and
carpopodium is not wrapped. In A. argyi,
A. dracunculus, and A. marschalliana, one-two
lower rows of carpopodial cells enter the inner
surface of the basal part of the achene, forming
a wrapped carpopodium. In A. absinthium,
A. annua, A. argyi, and A. marschalliana, the
primary ultrastructure of the carpopodium is
well defined, and cells have clear boundaries.
In A. dracunculus and A. vulgaris, the primary
ultrastructure is fuzzy; the cells’ contours are
visible only in the upper rows; they are smooth
in the rows below. Straightened smooth cuticle
characterizes the secondary ultrastructure of
the carpopodium.
The place of attachment of the corolla at the
apex of the achene (apical bowl) is well defined
in all species. It is outlined by a rim, which is
placed in the middle of the apex or is slightly
shifted to the side due to the topography
of the fruit (Boyko, 2011). The presence and
nature of the rim are strictly related to the
embryo size. The rim appears due to the
embryo’s folding due to drying of the achene
(Korobkov, 1973). In A. absinthium, the rim is
lobed, rounded, formed by the anticlinal walls
of mucous cells. In other investigated species,
the rim is solid, rounded, symmetrical, formed
by the anticlinal walls of both mucous and
epidermal cells of the exocarp. In A. vulgaris
the rim is the highest and has the shape of a
thin membranous crown. In A. dracunculus
the place of attachment of a corolla is well
defined, but the rim is absent. The rim with
smallest diameter is typical to A. marschalliana
(up to 90 μm) and A. annua (up to 100 μm); in
other species, it is almost twice as wide (220–
250 μm).
Specialized mucous cells were found in
the exocarp of the studied species. Among
Asteraceae, such mucous cell complexes
were found only in the achenes of the
representatives of the tribe Anthemideae.
These complexes serve as a reliable taxonomic
sign of this tribe but do not distinguish genera
(Yakovleva et al., 2002; Mungalov, 2004; Boyko,
2010). Mucous cells are usually grouped into
mucous complexes in the form of longitudinal
spherical cords extending from the apex to
the base of the achene. These cords are raised
above the achene’s surface in A. absinthium,
A. argyi, and A. marschalliana. However, in
A. annua, A. dracunculus, and A. vulgaris, they
are located almost on the exocarp surface
level. In the achenes of A. vulgaris that we
studied, the cords were poorly visible; the
mucous cells were located mainly in separate
groups. Although some literature data indicate
long mucous cords in A. vulgaris, other sources
reported their absence (Korobkov, 1973; Boyko,
2013). This discrepancy could be explained
by the phenomenon when in some Artemisia
species, the mucous cells lose their functions
but retain their characteristic structure.
The ribbed surface of the achenes of
Artemisia can have a different origin. The
SEM made it possible to clarify the nature of
the microrelief of studied species’ achenes.
Many authors (Korobkov, 1973; Beer, 2005;
Boyko, 2011) distinguish between longitudinal
streaking of achenes (due to the presence
in the epidermis of the exocarp of large
mucous complexes represented by strands
longitudinally raised above the body of the
achene) and longitudinal wrinkles, which are
loosely adjacent to the achene. Among the
studied achenes, the longitudinally striped
surface we found in A. absinthium and
A. marschalliana. At the same time, achenes
of A. argyi, A. dracunculus, and A. vulgaris are
characterized by a longitudinally wrinkled
surface. In A. annua achenes, the nature
of the surface is somewhat different from
other species: the mucous membranes are
quite broad; the main cells of the exocarp
are arranged between them in a row and
significantly compressed. The mucous cells’
outer periclinal walls are concave or flat, and
wide rounded branched folds clearly appear
through them, forming a kind of reticular
pattern. This appearance of the achene’s
surface with thin embryo is not caused by
the structure of the exocarp solely. Also, it is
affected by the relief of the exotests placed
under it, like in many asters with a tight fit of
the embryo to the test (Boyko & Novozhilova,
2018). We defined the type of surface found in
A. annua as reticulate-pitted.
On the surface of the achenes of A. argyi,
A. dracunculus, and A. vulgaris schizogenic
secretory containers with well-marked
polygonally rounded squamous cells were
found. The primary ultrastructure of mucous
cells and regular exocarp cells located
Plant Introduction • 87/88 63
Carpological features of certain Artemisia species distributed in Ukraine
between the mucous cords is somewhat
different. Mucous cells are the most well-
defined. In all investigated species, they have
relatively clear boundaries; their anticlinal
walls are straight (A. absinthium, A. argyi, and
A. dracunculus), or slightly wavy, unevenly
thickened (A. annua and A. vulgaris). The
outer periclinal walls are usually concave or
flat, sometimes with a tubercle in the middle
(A. argyi and A. marschalliana) that protrudes
through the outer wall. Such bumps may
indicate the presence in exocarp cells of
various crystalline inclusions or individual
large crystals of calcium oxalate, which is
characteristic of many wormwoods (Boyko,
2014). The primary ultrastructure of epidermal
cells of the exocarp is weakly defined. These
cells are significantly compressed, often
without clear boundaries, with unevenly
thickened anticlinal walls, the edges of which
are straight (A. argyi and A. dracunculus),
wavy (A. absinthium and A. marschalliana), or
fine-toothed (A. annua). The outer periclinal
wall is concave, rarely flat, or slightly convex;
its surface is often wrinkled or folded. It is
proved that such folding is formed not by
the cuticle but by the irregularities of the
outermost periclinal walls (Yakovleva et al.,
2002). Secondary ultrastructure is defined
differently in different Artemisia species. In
particular, A. absinthium is characterized by
a pronounced folded cuticle over the mucous
membranes. It was found that the thickened
cuticle characterizes the secondary structure
of A. marschaliana. Sometimes such thickened
layers form cuticular conglomerates. In
other species (A. absinthium, A. annua, and
A. vulgaris), the cuticle is more or less straight;
cuticular tubercles and growths are only
occasionally present on its surface.
Conclusions
As a result of comparative morphological
analysis of achenes of six species of the genus
Artemisia, a specific variety of individual
structural elements was revealed, the features
of which can be used comprehensively for
reliable identification of species. These
features can be used to identify investigated
Artemisia species by fruit, especially when their
recognition based on macromorphological
features is complicated.
The set of diagnostic carpological criteria to
distinguish Artemisia species should include:
a) such peculiarities of the carpopodium as
nature of the line of demarcation between
the carpopodium and the achene body, and
location of carpopodial cells on the inner
surface of the basal part of the achene;
b) presence or absence of apical rim; c) features
of primary and secondary ultrastructure of the
exocarp’s strands and epidermal cells.
References
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Карпологічні особливості деяких представників роду Artemisіa, поширених в
Україні
Тетяна Вакуленко *, Тетяна Багацька, Ольга Корабльова **, Іван Гурненко, Джамал Рахметов
Національний ботанічний сад імені М.М. Гришка НАН України, вул. Тимірязєвська, 1, Київ, 01014,
Україна; * botanicukr@gmail.com, ** okorablova@ukr.net
Досліджено мікроморфологічну будову сім’янок шести видів роду Artemisia (A. absinthium, A. annua,
A. argyi, A. dracunculus, A. marschalliana та A. vulgaris), представлених у флорі України, з метою виявити
карпологічні ознаки, які можуть слугувати діагностичними для визначенні цих видів.
Плоди збирали з дикорослих рослин рекреаційних зон міста Києва, та тих, що зростають
на колекційній ділянці Національного ботанічного саду імені М.М. Гришка НАН України.
Макроморфологічні особливості вивчали за допомогою бінокулярного мікроскопа Stemi 2000,
Carl Zeiss. Ультраструктуру поверхні вивчали за допомогою растрового електронного мікроскопа
JSM-6700F, JEOL у режимі вторинної електронної емісії. Напилення вуглецем здійснювали методом
вакуумного термічного випаровування вакуумного універсального поста ВУП-5М, SELMI; напилення
платиною – за допомогою пристрою іонно-плазмового травлення JFC-1600, JEOL.
Усі досліджені види мають добре виражений, справжній, багатоклітинний, симетричний,
багаторядний карпоподій, який, однак, відрізняється за формою. Тип поверхні у A. annua визначено
як сітчасто-ямковий, що зумовлено структурою екзокарпа та екзотестами, розташованими під
ним. Виявлено види з лопатевим або суцільним верхівковим обідком, а також такі, у яких обідок
навколо місця прикріплення віночка відсутній. Встановлено, що первинна ультраструктура
плодів спричинена формою та характером розташування слизових клітин; епідермальні клітини
екзокарпа переважно стиснуті, без чітких меж. В той час як вторинна ультраструктура плодів
зумовлена переважно тонкою і непомітною кутикулою, іноді зморшкуватою, з прошарками і
невеликими наростами. Досліджені карпологічні ознаки можуть бути додатковими критеріями та
використовуватись у комплексі з іншими ознаками для точнішої ідентифікації досліджених видів.
Ключові слова: Asteraceae, Artemisia, карпоподіум, слизові комплекси, первинна ультраструктура, вторинна
ультраструктура, плід
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| id | oai:ojs2.plantintroduction.org:article-1572 |
| institution | Plant Introduction |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2025-07-17T12:53:47Z |
| publishDate | 2020 |
| publisher | M.M. Gryshko National Botanical Garden of the NAS of Ukraine |
| record_format | ojs |
| resource_txt_mv | wwwplantintroductionorg/f6/3c69fe18fc1494fb77c41af06d5c69f6.pdf |
| spelling | oai:ojs2.plantintroduction.org:article-15722023-08-26T20:39:33Z Carpological features of certain Artemisia species distributed in Ukraine Карпологічні особливості деяких представників роду Artemisіa, поширених в Україні Vakulenko, Tetyana Bagatska, Tetyana Korablova, Olga Gurnenko, Ivan Rakhmetov, Dzhamal The achenes’ micromorphological structure of six Artemisia species (A. absinthium, A. annua, A. argyi, A. dracunculus, A. marschalliana, and A. vulgaris) distributed in Ukraine was investigated with the aim to identify taxonomically significant carpological features for the studied species.Fruits were collected from wild plants growing in the recreation areas of Kyiv and from the plants hosted in the living collection of the M.M. Gryshko National Botanical Garden of the NAS of Ukraine. Macromorphological features were studied with a binocular microscope Stemi 2000 Carl Zeiss. The ultrastructure of the surface was studied using a scanning electron microscope JSM-6700F JEOL, in the mode of secondary electron emission. Carbon spraying was performed using vacuum thermal evaporation of the vacuum universal post VUP-5M SELMI; platinum spraying was performed with an ion-plasma etching device JFC-1600 JEOL.All investigated species have a well-expressed, true, multicellular, symmetrical, multi-row carpopodium but differ by its shape. The surface of A. annua achenes is reticulate-pitted due to the specific structure of the exocarp and the exotests located below it. Species with lobed and continuous apical rim and species without a rim around the place of attachment of the corolla were found. It was established that the primary ultrastructure of investigated fruits is caused by the shape and character of mucous cell arrangement; epidermal cells of the exocarp are mostly compressed, without clear boundaries. Mostly a thin and inconspicuous cuticle, sometimes wrinkled, with layers and small outgrowths, causes the secondary ultrastructure of these fruits. We believe that the mentioned carpological features may be used as additional criteria in combination with other features to identify studied species. Досліджено мікроморфологічну будову сім’янок шести видів роду Artemisia (A. absinthium, A. annua, A. argyi, A. dracunculus, A. marschalliana та A. vulgaris), представлених у флорі України, з метою виявити карпологічні ознаки, які можуть слугувати діагностичними для визначенні цих видів.Плоди збирали з дикорослих рослин рекреаційних зон міста Києва, та тих, що зростають на колекційній ділянці Національного ботанічного саду імені М.М. Гришка НАН України. Макроморфологічні особливості вивчали за допомогою бінокулярного мікроскопа Stemi 2000, Carl Zeiss. Ультраструктуру поверхні вивчали за допомогою растрового електронного мікроскопа JSM-6700F, JEOL у режимі вторинної електронної емісії. Напилення вуглецем здійснювали методом вакуумного термічного випаровування вакуумного універсального поста ВУП-5М, SELMI; напилення платиною – за допомогою пристрою іонно-плазмового травлення JFC-1600, JEOL.Усі досліджені види мають добре виражений, справжній, багатоклітинний, симетричний, багаторядний карпоподій, який, однак, відрізняється за формою. Тип поверхні у A. annua визначено як сітчасто-ямковий, що зумовлено структурою екзокарпа та екзотестами, розташованими під ним. Виявлено види з лопатевим або суцільним верхівковим обідком, а також такі, у яких обідок навколо місця прикріплення віночка відсутній. Встановлено, що первинна ультраструктура плодів спричинена формою та характером розташування слизових клітин; епідермальні клітини екзокарпа переважно стиснуті, без чітких меж. В той час як вторинна ультраструктура плодів зумовлена переважно тонкою і непомітною кутикулою, іноді зморшкуватою, з прошарками і невеликими наростами. Досліджені карпологічні ознаки можуть бути додатковими критеріями та використовуватись у комплексі з іншими ознаками для точнішої ідентифікації досліджених видів. M.M. Gryshko National Botanical Garden of the NAS of Ukraine 2020-12-30 Article Article application/pdf https://www.plantintroduction.org/index.php/pi/article/view/1572 10.46341/PI2020038 Plant Introduction; No 87/88 (2020); 54-64 Інтродукція Рослин; № 87/88 (2020); 54-64 2663-290X 1605-6574 10.46341/PI87-88 en https://www.plantintroduction.org/index.php/pi/article/view/1572/1499 Copyright (c) 2020 Tetyana Vakulenko, Tetyana Bagatska, Olga Korablova, Ivan Gurnenko, Dzhamal Rakhmetov http://creativecommons.org/licenses/by/4.0 |
| spellingShingle | Vakulenko, Tetyana Bagatska, Tetyana Korablova, Olga Gurnenko, Ivan Rakhmetov, Dzhamal Карпологічні особливості деяких представників роду Artemisіa, поширених в Україні |
| title | Карпологічні особливості деяких представників роду Artemisіa, поширених в Україні |
| title_alt | Carpological features of certain Artemisia species distributed in Ukraine |
| title_full | Карпологічні особливості деяких представників роду Artemisіa, поширених в Україні |
| title_fullStr | Карпологічні особливості деяких представників роду Artemisіa, поширених в Україні |
| title_full_unstemmed | Карпологічні особливості деяких представників роду Artemisіa, поширених в Україні |
| title_short | Карпологічні особливості деяких представників роду Artemisіa, поширених в Україні |
| title_sort | карпологічні особливості деяких представників роду artemisіa, поширених в україні |
| url | https://www.plantintroduction.org/index.php/pi/article/view/1572 |
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