Мікроморфологія та анатомія плоду Iris pseudacorus L.
Based on the suggestion that flower and fruit are integrally evolving structures, we aimed to reveal the floral traits persisting in the fruit structure in Iris pseudacorus, a widely distributed riparian species in Ukraine. We intended to compare the results with the other Iris species studied previ...
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| Дата: | 2023 |
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M.M. Gryshko National Botanical Garden of the NAS of Ukraine
2023
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Plant Introduction| _version_ | 1860145144213274624 |
|---|---|
| author | Odintsova, Anastasiya Khomei, Yaroslav |
| author_facet | Odintsova, Anastasiya Khomei, Yaroslav |
| author_sort | Odintsova, Anastasiya |
| baseUrl_str | https://www.plantintroduction.org/index.php/pi/oai |
| collection | OJS |
| datestamp_date | 2024-04-07T19:57:15Z |
| description | Based on the suggestion that flower and fruit are integrally evolving structures, we aimed to reveal the floral traits persisting in the fruit structure in Iris pseudacorus, a widely distributed riparian species in Ukraine. We intended to compare the results with the other Iris species studied previously and reveal the constancy of micromorphological features of fruit interior structure. We revealed exomorphological and micromorphological peculiarities of the fruiting ovary using the model of vertical zonality of the gynoecium, vascular anatomy, and general anatomy of the fruit wall. In the fruiting ovary of I. pseudacorus, we revealed the presence of three vertical zones: short synascidiate zone, long symplicate zone bearing uniseriate seeds, and hemisymplicate zone located in the fruit beak. The vascular system of the ovary is composed of dorsal, septal, and ventral veins. Each of three dorsal veins divides radially into the outer tepal trace, stamen trace, and dorsal carpellary bundle, while each septal vein divides tangentially into three bundles of the inner tepal trace. Paired ventral veins enter the ovary from its bottom and supply ovules and seeds. The exocarp is composed of polygonal cells with thickened cellulose walls. The endocarp is uniseriate, unlignified, and composed of live prosenchymal cells, which are elongated tangentially. In the parenchymatous mesocarp, a great number of secretory canals with tannin-like content occur. The dehiscence of fruit on three valves proceeded by both dorsal and ventral slits. Dorsal slits are formed along dorsal grooves and provided by small-celled tissue surrounding the dorsal veins. The presence of ventral sutures of carpels in the symplicate zone of the ovary provides ventral dehiscence of fruit. Hence, the structure of the fruiting ovary in I. pseudacorus is comparable to that of other Iris species. Our investigation confirmed that the vertical zonality, placentation, and vascular system of the gynoecium in Iris can be appropriately estimated in the fruiting stage because the structural components of the ovary, which developed at the pre-anthetic phase, persist in the fruit. |
| doi_str_mv | 10.46341/PI2023004 |
| first_indexed | 2025-07-17T12:54:16Z |
| format | Article |
| fulltext |
© The Authors. This content is provided under CC BY 4.0 license.
Plant Introduction, 99/100, 12–23 (2023)
RESEARCH ARTICLE
Micromorphology and anatomy of fruit in Iris pseudacorus L.
Anastasiya Odintsova *, Yaroslav Khomei
Ivan Franko National University of Lviv, Hrushevskyi str. 4, 79005 Lviv, Ukraine; * anastasiya.odintsova@lnu.edu.ua
Received: 27.07.2023 | Accepted: 04.09.2023 | Published online: 24.09.2023
Abstract
Based on the suggestion that flower and fruit are integrally evolving structures, we aimed to
reveal the floral traits persisting in the fruit structure in Iris pseudacorus, a widely distributed
riparian species in Ukraine. We intended to compare the results with the other Iris species studied
previously and reveal the constancy of micromorphological features of fruit interior structure.
We revealed exomorphological and micromorphological peculiarities of the fruiting ovary using
the model of vertical zonality of the gynoecium, vascular anatomy, and general anatomy of the
fruit wall. In the fruiting ovary of I. pseudacorus, we revealed the presence of three vertical zones:
short synascidiate zone, long symplicate zone bearing uniseriate seeds, and hemisymplicate zone
located in the fruit beak. The vascular system of the ovary is composed of dorsal, septal, and
ventral veins. Each of three dorsal veins divides radially into the outer tepal trace, stamen trace,
and dorsal carpellary bundle, while each septal vein divides tangentially into three bundles of
the inner tepal trace. Paired ventral veins enter the ovary from its bottom and supply ovules and
seeds. The exocarp is composed of polygonal cells with thickened cellulose walls. The endocarp is
uniseriate, unlignified, and composed of live prosenchymal cells, which are elongated tangentially.
In the parenchymatous mesocarp, a great number of secretory canals with tannin-like content
occur. The dehiscence of fruit on three valves proceeded by both dorsal and ventral slits. Dorsal
slits are formed along dorsal grooves and provided by small-celled tissue surrounding the dorsal
veins. The presence of ventral sutures of carpels in the symplicate zone of the ovary provides
ventral dehiscence of fruit. Hence, the structure of the fruiting ovary in I. pseudacorus is comparable
to that of other Iris species. Our investigation confirmed that the vertical zonality, placentation,
and vascular system of the gynoecium in Iris can be appropriately estimated in the fruiting stage
because the structural components of the ovary, which developed at the pre-anthetic phase,
persist in the fruit.
Keywords: Iris, vascular anatomy, symplicate zone, fruit wall, fruiting ovary
https://doi.org/10.46341/PI2023004
UDC 581.47 : 582.579.2
Authors’ contributions: Both authors sampled plant material, conducted microscopic studies, described the results, and wrote
the text of the article. Anastasiya Odintsova made general supervision of the article, adopted the methodology, and wrote
the original draft of the manuscript. Yaroslav Khomei provided microtechnique procedures, sectioned the material, made
microphotographs, and designed the text of the article.
Funding: The study was conducted within the unfunded project: “Structural diversity and morphogenesis of reproductive organs of
angiosperms at individual and population levels” (state registration Nr 0122U200558).
Competing Interests: The authors declare that they have no conflict of interest.
https://creativecommons.org/licenses/by/4.0/
https://orcid.org/0000-0001-7645-3311
Plant Introduction • 99/100 13
Micromorphology and anatomy of fruit in Iris pseudacorus L.
Introduction
Iris L. is the most species-rich genus in the
family Iridaceae, being treated morphologically
for taxonomic purposes, mainly from the
flower and vegetative morphology (Fomin
& Bordzilovskyi, 1950; Goldblatt et al., 1998;
Wilson, 2006; Sennikov et al., 2023). However,
fruit structure in this genus, especially with
the application of anatomical techniques, was
poorly studied (Rodionenko, 1961; Gritsenko,
2020; Skrypec & Odintsova, 2020). For the
whole Iridaceae family, the fruit was defined as
a syncarpous inferior trilocular multi-seeded
capsule with loculicidal dehiscence (Goldblatt
et al., 1998). Two distinct dehiscence types
were described for different Iris species
(Rodionenko, 1961). The first type of fruit
dehiscence is realized through the separation
of the fruit into three declined valves formed
by dorsal slits going through the whole fruit or
only through its upper part. In some species,
a column persists in the center of the fruit
after valves are declined. The second type of
fruit dehiscence proceeds by three dorsal
slits without forming valves, so the fruit does
not disintegrate. Since we suggest flower and
fruit as a united evolving structure (Odintsova,
2022), we aimed to reveal the floral traits
persisting in the fruit structure in Iris species
that have large fleshy fruits suitable for
anatomical research.
Being experienced in the study of flower
and fruit morphology and anatomy in Iris
sibirica L. (Skrypec & Odintsova, 2014, 2015,
2020), we intended to compare previous
results with data on the I. pseudacorus L., a
species belonging to the same section Limniris
of the subgenus Limniris. The objectives of the
present study were to reveal the constancy of
micromorphological features of fruit interior
structure, including those of the gynoecium
(vertical zonality, placentation, vascular
anatomy). The anatomy of the fruit wall was
also investigated as having a substantial role in
fruit dehiscence.
Iris pseudacorus is an Eurasian riparian
species, growing in Ukraine throughout the
entire country except wormwood steppe and
Crimea (Zhygalova, 2014). It belongs to the series
Laevigatae of the section Limniris (Rodionenko,
1961; Wilson, 2009). The habitats of I. pseudacorus
are limited to the river banks, swamps, and
ponds. It is highly tolerant to submersion, water
pH, and soil properties, and is well reproduced
by rhizome and water-dispersed seeds. The
species has LC conservation status (IUCN,
2022) and is invasive in North America, where
it escapes from cultivation spots (Gerwing
et al., 2021; Stoneburner et al., 2021). Currently,
I. pseudacorus is cultivated as an ornamental
plant, while previously, its rhizome, being rich
in tannins, was used to obtain a black color after
adding ferrum salts, and flowers treated with
acetic acid were used to get a yellow color for skin
staining (Fomin & Bordzilovskyi, 1950). Tannins
are abundant in the vegetative structures and
flowers of some Iris species (Rodionenko, 1961).
In I. pseudacorus, the phenolic compounds
extracted from the rhizome were approved for
antibacterial activity (Michalak et al., 2021).
Material and methods
The unripe green fruits of Iris pseudacorus L.
were collected near Ivano-Frankove village,
Yavoriv district, Lviv oblast (Ukraine), in the
pond and channels (22 June 2015; 24 and 30
June 2023). Sampled materials were sectioned
by hand in fresh condition and after fixation in
70 % ethanol. Obtained sections were treated
with safranin solution or phloroglucinol
reaction with chloride acid for lignin (Gaertner
& Schweingruber, 2013; Pradhan Mitra &
Loqué, 2014). Twenty fruits were examined in
total. Images were captured by digital camera
Sigeta M3CMOS 10000 applied to the light
microscope XS-2610 (China).
For the micromorphological treatment,
we used Leinfellner’s (1950) model of vertical
zonality of the gynoecium. For vascular
anatomy, we followed Esau (1977) principles.
The anatomy of the fruit wall was surveyed
according to Roth (1977). As fruits of Iris are
arranged in monochasial cyme inflorescence,
they were at different stages of development
at the sampling time. Due to this reason, we
withheld quantitative data and statistical
evaluation of the studied features.
Results
Fruit morphology and micromorphology
Fruits of I. pseudacorus at the studied stages
were 5–8 cm long, banana-like, with a short
apical beak (spout), obtuse-triangular in a
14 Plant Introduction • 99/100
Odintsova & Khomei
transversal section, green and fleshy. Dorsal
grooves were well defined (Fig. 1 A). In our
materials, fruits had various sizes and
shapes; sometimes, they were deformed and
narrowed, depending on the abundance of
seeds inside (Fig. 1 A–C). On the transverse
section of the fruit, uniseriate pleurotropous
seed arrangement in the locule was evident
(Fig. 1 D). Many seeds in the fruit were
underdeveloped (Fig. 1 Е), and sometimes
one locule was sterile (Fig. 1 D). Seeds were
located along the whole locule height, from
the very bottom to its top; in the fruit with
low seed numbers, they occupy mostly the
upper part of the locule (Fig. 1 F). On the
radial section, it was visible that the fruit
beak is solid (locules do not extend in it),
and the abscission zone of the floral tube
lies beneath the style base (Fig. 1 F), which
means that the separation layer is formed in
the beak tissues where floral tube and style
are still fused.
On the transverse sections treated with
safranin, a short zone with congenitally closed
locules (without ventral sutures) was visible
(Fig. 2 A). In this zone, locules were narrow, and
seeds missing. Above it, ventral slits of carpels
appeared from the center and the locule side,
so a trilocular zone with a triradiate slit in the
center was formed. This slit connected locules
by the secretory epidermis (transmitting tract
to the ovules in the anthetic ovary) (Fig. 2 B–E).
Sometimes, at the base of this zone, septas
did not touch each other, and a small gap was
formed (Fig. 2 E). Seeds were born alternately
at one of each carpel’s margins. At the upper
portion of the ovary, locules were diminished,
and style channels formed from their proximal
parts, covered with secretory epidermis
(Fig. 2 F). In the fruit beak, initially triradiate
style channel became labyrinthine, and then,
the carpelar margins became free, so another
three slits arose between style channels, and the
central slit became hexaradiate (Fig. 3 А–D).
BA C F
D E
Figure 1. Exomorphology and interior structure of Iris pseudacorus fruit: A–C – unripe fruits of different
shapes; D – transversely sectioned fruit with one sterile and two fertile locules with uniseriate seed
arrangement; E – fragment of the longitudinally sectioned fruit with developing seeds and depressed
ovules (arrows); F – longitudinal section of the whole fruit (fb – fruit beak). A–D – fresh material; E, F – fixed
material.
Plant Introduction • 99/100 15
Micromorphology and anatomy of fruit in Iris pseudacorus L.
A B
C D
E F
Figure 2. Micromorphology and anatomy of Iris pseudacorus fruit body on transversal sections:
A – synascidiate zone of the fruiting gynoecium; B–D – transitions from synascidiate to symplicate zone,
ventral suture marked in D with black arrows; E – symplicate zone with a gap in the center between
incomplete septas; F – roof of the inferior ovary. dc – dorsal carpellary bundle; lo – locule; ov – ovule/
seed; tt – transmitting tissue; vc – ventral carpellary veins; vs – ventral suture, small vascular bundles are
marked with white arrows. A–D – safranin staining.
Therefore, in the fruiting ovary of
I. pseudacorus, we revealed the presence of
three vertical zones (Fig. 4): short and sterile
synascidiate zone, long symplicate zone
bearing placentas on the incomplete septas,
and hemisymplicate zone in the fruit beak.
The last zone continued into the style.
16 Plant Introduction • 99/100
Odintsova & Khomei
Vascular anatomy
On the series of transversal sections through
the fruit, we can trace the arrangement of
vascular bundles in the ovary. In the upper
portion of I. pseudacorus peduncle, a dicyclic
atactostele occurred, with an external ring
composed of numerous closely spaced
vascular bundles of various sizes and the
inner ring composed of six bundles located
on the tepal radii (Fig. 5 А). Above it, the
inner bundles entered the external ring, the
last divided into six trunk bundles entering
the fruit wall, and the rest of the vascular
bundles in the center of the ovary (Fig. 5
B, С). Three trunk bundles on the outer
tepal’s radii (on the carpel median plane)
were doubled (Fig. 5 B), – we entitled them
dorsal veins of carpels. The other three
trunk bundles located on the septas radii
were entitled septal veins. The remnants
of the vascular tissue formed a plexus of
small bundles, which develop paired ventral
carpellary bundles in the ends of septas.
These bundles gave rise to ovule traces
(Fig. 2 А–С). Small anastomoses were formed
between septal veins and ventral bundles,
especially in the lower portion of the fruit
(Figs. 2 В–D; 5 D, E). Dorsal veins gave tiny
branches to the ovary wall (Fig. 5 F).
In the fruiting ovary, dorsal and septal
veins were unitary (Fig. 5 Е, F), while in the
fruit beak, each dorsal vein branched radially
onto three bundles: outer tepal trace, stamen
Figure 3. Micromorphology and anatomy of Iris pseudacorus fruit beak on ascending transversal sections
(A→D): dc – dorsal carpellary bundle; dv – dorsal vein; itt – inner tepal trace; ott – outer tepal trace; sc – style
canals; st – stamen trace; tt – transmitting tissue; vv – ventral veins; arrows in B mark transmitting tissue
in hemisymplicate zone. A, C, D – safranin staining.
A B
DC
Plant Introduction • 99/100 17
Micromorphology and anatomy of fruit in Iris pseudacorus L.
Figure 4. Schematic drawing of the fruit zonality and vascularization: Hspl zone – hemisymplicate zone; Sa
zone – synascidiate zone; Spl zone – symplicate zone. Postgenitally fused epidermises of ventral sutures
and carpel margins are colored in grey. dc – dorsal carpellary bundle; dv – dorsal vein; itt – inner tepal
trace; ott – outer tepal trace; st – staminal trace; sv – septal vein; tt – transmitting tissue; vv – ventral veins.
trace, and dorsal carpellary bundle, which
declined to the center and entered the style
(Fig. 3 А). Stamen trace was basally concentric
with xylem outwards (Fig. 3 A). The septal vein
in the fruit beak divided into three bundles
tangentially: two lateral bundles were larger
than the central one and made tiny branches
to the center (Fig. 3 A, С). Ventral bundles
in the beak fused in each septa ending
(Figs. 3 С; 4).
Therefore, the vascular system of
I. pseudacorus ovary can be appropriately
estimated during the fruiting stage because
vascular bundles, which mainly developed
at the pre-anthetic phase, persist in the
fruit. The vascular system of the epigynous
flower of I. pseudacorus is integrated: dorsal
carpellary bundles are condensed with tepal
and stamen traces, while septal veins are
anastomosed with ventral carpellary bundles.
Six trunk bundles supply the fruit wall, while
ovules/seeds are supplied from ventral
bundles entering the ovary from its bottom.
In the beak, which is the narrowest part of
the fruit, trunk bundles branch out. Small
bundles between all main veins are visible.
The occurrence of many small bundles in the
fruit wall and septas is a regular characteristic
of large fleshy fruits (Roth, 1977).
Anatomy of the fruit wall
The fruit wall of I. pseudacorus at the stage of
green fruit was parenchymatous and fleshy.
The exocarp was uniseriate, composed of
polygonal isodiametric cells with thickened
cellulose walls and a thick cuticle on the
outer wall (Fig. 3 С). Stomata in the exocarp
were surrounded by 4–7 cells, not sunken
(Fig. 6 A, C). Endocarp at the investigated
stage of fruit development was uniseriate,
unlignified, and composed of prosenchyma
cells, elongated tangentially, 200–400
mkm long, with oblique ends (Fig. 6 B, D).
Endocarpial cells had evenly thickened walls
and remained alive.
The mesocarp was at least 1.7 mm thick
and composed of 27–38 cell layers (in the
18 Plant Introduction • 99/100
Odintsova & Khomei
A
C
E
B
D
F
Figure 5. The vascular system of Iris pseudacorus fruit on transversal sections: A – the base of the fruiting
ovary; B – formation of dorsal veins in receptacle; C – formation of septal and ventral veins at the locule
base; D, E – septal vein at the middle height of the fruiting ovary, anastomosing to ventral veins; F – dorsal
vein. ab – anastomosing bundle; dv – dorsal vein; lo – locule; scp – small-celled parenchyma; sv – septal
vein. A – phloroglucinol-HCl staining; B, D–F – safranin staining.
median plane of the carpel near the dorsal
vein). Mesocarp cells had almost equal
sizes, but external chlorenchyma cells were
slightly smaller. Chlorenchyma appeared
in the external zone of the mesocarp as a
continual layer; the inner zone of the fruit
wall was colorless (Fig. 1 D). In the mesocarp
of I. pseudacorus, the sporadic appearance of
crystallin styloids, up to about 155 mkm long,
was noticed (Fig. 7 A, B). Many secretory
scp
Plant Introduction • 99/100 19
Micromorphology and anatomy of fruit in Iris pseudacorus L.
sc
canals occured in mesocarp, arranged
solitary or grouped (Fig. 7 C, D). The secretory
canals were 0.4–2.8 mm long (Fig. 7 C). They
appeared in the external and inner zones of
the mesocarp, in septas, in the fruit base, and
a beak (Figs. 2; 3; 5; 7D). On fresh untreated
sections of the fruit, they looked like round
cavities, larger than parenchyma cells, while
after safranin treatment, they showed a red
outline or filling as tannins do. In the fixed
material, the canals had yellow content
(Fig. 7 C, D).
A significant element of the fruit
dehiscence is the arrangement of
the mechanical tissues enabling slit
development. Morphologically, dorsal slits
are predicted by grooves on the external
face of the fruit. Anatomically, on the median
plane of the carpel, proximally to the dorsal
vein, small uncolored live cells appear in the
mesocarp (Fig. 5 F; 7 D). Such small cells are
precursors of dorsal slits of the fruit in many
capsular fruits (Roth, 1977). The dehiscence
of the capsule in I. pseudacorus was
designated by Rodionenko (1961) as valvate.
Three declined valves of fruit were evident
in the photos of fruit made by Kovalchuk
(2009); the dehiscing fruits on the images
have a greenish color, and they are not dry
and brown when dehisce. The disintegration
of fruit on the valves proceeded by dorsal
and ventral slits through the whole fruit.
The presence of ventral sutures along the
fruiting ovary supports ventral dehiscence in
this species.
Discussion
Our research revealed the fruit characteristics
common for I. pseudacorus and I. sibirica. In
both species, the size of the fruit depends on
the number of seeds inside; seeds are located
horizontally from the very base to the top of
A
C
B
D
Figure 6. Epidermises of the fruit of Iris pseudacorus: A, С – exocarpium with stomata; B, D – endocarpium;
A, B – paradermal sections; C, D – transversal sections of the fruit. sc – secretory canal.
20 Plant Introduction • 99/100
Odintsova & Khomei
scp
A
C
B
D
Figure 7. Anatomy of fruit wall in Iris pseudacorus: A, B – cristallic inclusion-styloid (arrow) in mesocarp;
C – secretory canals on the tangential section and D – on the transverse section in the dorsal portion of
the carpel. Dorsal vein (dv) and small-celled parenchyma (scp) are visible along the upcoming dorsal slit
toward the locule (lo).
the locule (Skrypec & Odintsova, 2015). After
anthesis, the floral tube falls out with an
accrescent style in both species (Skrypec &
Odintsova, 2020). However, in I. pseudacorus
seeds arrangement is uniseriate, contrary to
the biseriate arrangement in I. sibirica and
many other Iris species (Rodionenko, 1961);
dorsal grooves are visible in the green fruit,
and slits are complete (arise from the top to
base of fruit) contrary to I. sibirica, where
grooves are not detectable and slits are mostly
short (Skrypec & Odintsova, 2014).
The interior structure of the fruiting
ovary in I. pseudacorus is characterized by
the appearance of three zones. However,
the synascidiate zone is short and sterile,
contrary to I. sibirica (Skrypec & Odintsova,
2020). The appearance of synascidiate and
symplicate zones in the ovary (but without
entitling these zones) was clearly illustrated
by Van Tieghem (1871) for I. lutescens Lam.
(mentioned as I. chamaeiris Bertol in the
original work). The sterile hemisymplicate
zone at the upper part of the ovary and style
is represented by Rodionenko (1961) for
I. pseudacorus, I. orientalis Mill. (mentioned
as I. monnieri DC. in the original work), and
I. korolkowii Regel in fig. 26, while synascidiate
and symplicate zones in some Iris species –
in figs. 45–48. Those data suggested defining
the gynoecium in Iris species and Iridасеае in
general as eusyncarpous gynoecium according
to Leinfellner’s (1950) classification. The
gynoecium having synascidiate, symplicate,
and hemisymplicate zones is a widely
distributed type of the gynoecium among
monocots (Novikoff & Odintsova, 2008);
however, the length and ratio of zones vary
in a wide range. For instance, hemisymplicate
zone can occupy a large portion of the ovary
when it contains septal nectaries, which are
missing in Iris.
Plant Introduction • 99/100 21
Micromorphology and anatomy of fruit in Iris pseudacorus L.
The vascular system of the ovary in
I. pseudacorus is very similar to the vascular
system of other species of Iridасеае studied
previously (Van Tieghem, 1871; Rodionenko,
1961; Skrypec & Odintsova, 2020). The main
feature of the vascularization of the ovary is
the occurrence of three types of veins: dorsal,
septal, and ventral. The common pattern of
the vascular system is a radial divide of the
dorsal vein into three traces (outer tepal trace,
stamen trace, and dorsal carpellary bundle),
and tangential divide of the septal vein into
three bundles, too. The exact structure of
the vascular system is usual for epigynous
flowers of lilioid monocots: i.e., Iris lutescens,
I. florentina L., Crocus vernus (L.) Hill.,
Gladiolus ×gandavensis Van Houtte, Narcissus
pseudonarcissus L. (Van Tieghem, 1871) as
well Iris sibirica and Gladiolus imbricаtus L.
(Skrypec & Odintsova, 2020).
The most peculiar feature of the fruit of
I. pseudacorus is the abundance of canals with
tannin-like content, which usually occur in
fruits (Roth, 1977). Considering the abundance
of the secretory canals in I. pseudacorus fruits,
we suggest using its fruits as medicinal raw
resources.
Many capsular fruits of lilioid monocots
have lignified endocarp, V-shaped xylem in the
dorsal vein, and U-shaped cell wall thickening
in the endocarp (Rasmussen et al., 2006).
Instead, some fleshy capsules or berry-like
fruits reveal delayed dehiscence and lacking
lignification in the fruit wall. They arise in
conditions of high humidity or in cases when
the capsule lies on the ground when ripe and
does not dehisce. Iridaceae was not mentioned
as a family with berry-like fruits (Rasmussen
et al., 2006; Thadeo et al., 2015), but we can
denote the capsule is a long-time fleshy,
lately desiccates and has delayed lignification
of the endocarp. This can be caused by the
hydrophilous habitats of the species.
Conclusions
The structure of the fruiting ovary in Iris
pseudacorus is comparable to the ovary of
other Iris species. Our investigation confirmed
that the vertical zonality, placentation, and
vascular system of the gynoecium in Iris could
be estimated correctly in the fruiting stage
because the structural components of the
ovary, which developed at the pre-anthetic
phase, persist in the fruit.
References
Esau, K. (1977). Anatomy of seed plants, 2nd ed. Wiley
and Sons.
Fomin, O.V., & Bordzilovskyi, Y.I. (1950). Family
Iridaceae Lindl. In: M.I. Kotov, A.I. Barbarych
(Eds.), Flora URSR, Vol. 3 (pp. 276–312). Academy
of Sciences of UkrSSR (In Ukrainian)
Gaertner, H., & Schweingruber, F.H. (2013).
Microscopic preparation techniques for plant
stem analysis. Dr. Kessel Verlag https://www.
researchgate.net/publication/253341899_
Microscopic_Preparation_Techniques_for_
Plant_Stem_Analysis
Gerwing, T.G., Thomson, H.M., Brouard-John, E.K.,
Kushneryk, K., Davies, M.M., Lawn, P., &
Nelson, K.R. (2021). Observed dispersal of
invasive yellow flag iris (Iris pseudacorus) through
a saline marine environment and growth in a
novel substrate, shell hash. Wetlands, 41, Article 1.
https://doi.org/10.1007/s13157-021-01421-w
Goldblatt, P., Manning, J.C., & Rudall, P. (1998).
Iridaceae. In: K. Kubitzki, H. Huber, P.J. Rudall,
P.S. Stevens & T. Stützel (Eds.), The families and
genera of vascular plants. III. Flowering plants:
Monocotyledons: Lilianae (except Orchidaceae) (pp.
295–333). Springer-Verlag.
Gritsenko, V.V. (2020). Morphological peculiarities
of fruits of the rare species Iris halophila Pall,
I. pumila L. and I. hungarica Waldst. et Kit.
(Iridaceae Juss.) in the conditions of introduction
in the meadow-steppe cultural phytocenosis.
Plant Introduction, 85/86, 85–92. https://doi.
org/10.46341/PI2020007
IUCN. (2022). The IUCN Red List of Threatened
Species. Version 2022-2. https://www.iucnredlist.
org/. Accessed on July 24, 2023.
Kovalchuk, A. (2009). Iris pseudacorus L. Image IDs
# 273797, 273798, 273799. In: UkrBIN: Ukrainian
Biodiversity Information Network (public project &
web application). https://ukrbin.com/show_image.
php?imageid=273797. Accessed on July 6, 2023.
Leinfellner, W. (1950). Der Bauplan des synkarpen
Gynözeums. Oesterreichische botanische Zeitschrift,
97, 403–436. https://doi.org/10.1007/BF01763317
Michalak, A., Krauze-Baranowska, M., Migas, P.,
Kawiak, A., Kokotkiewicz, A., & Królicka, A. (2021).
Iris pseudacorus as an easily accessible source of
antibacterial and cytotoxic compounds. Journal of
Pharmaceutical and Biomedical Analysis, 195, Article
113863, https://doi.org/10.1016/j.jpba.2020.113863
https://www.researchgate.net/publication/253341899_Microscopic_Preparation_Techniques_for_Plant_Stem
https://www.researchgate.net/publication/253341899_Microscopic_Preparation_Techniques_for_Plant_Stem
https://www.researchgate.net/publication/253341899_Microscopic_Preparation_Techniques_for_Plant_Stem
https://www.researchgate.net/publication/253341899_Microscopic_Preparation_Techniques_for_Plant_Stem
https://doi.org/10.1007/s13157-021-01421-w
https://doi.org/10.46341/PI2020007
https://doi.org/10.46341/PI2020007
https://www.iucnredlist.org/
https://www.iucnredlist.org/
https://ukrbin.com/show_image.php?imageid=273797
https://ukrbin.com/show_image.php?imageid=273797
https://doi.org/10.1007/BF01763317
https://doi.org/10.1016/j.jpba.2020.113863
22 Plant Introduction • 99/100
Odintsova & Khomei
Novikoff, A.V. & Odintsova, А. (2008). Some aspects
of gynoecium morphology in three bromeliad
species. Wulfenia, 15, 13–24.
Odintsova, A.V. (2022). Morphogenesis of fruit as
a subject matter for the carpological studies.
Ukrainian Botanical Journal, 79(3), 169–183.
(In Ukrainian). https://doi.org/10.15407/
ukrbotj79.03.169
Pradhan Mitra, P., & Loqué, D. (2014).
Histochemical staining of Arabidopsis thaliana
secondary cell wall elements. Journal of Visualized
Experiments: JoVE, 87, Article e51381. https://doi.
org/10.3791/51381
Rasmussen, F.N., Frederiksen, S., Johansen, B.,
Jorgensen, L.B., Petersen, G., & Seberg, O.
(2006). Fleshy fruits in liliiflorous monocots. Aliso:
A Journal of Systematic and Evolutionary Botany,
22(1), 135–147.
Rodionenko, G.I. (1961). Genus iris – Iris L.: questions
of morphology, biology, evolution and systematics.
Publishing House of the Academy of Sciences of
the USSR. (In Russian)
Roth, I. (1977). Fruits of angiosperms. In: W.
Zimmermann, S. Carlquist, P. Ozenda, & H.D.
Wulff (Eds.), Encyclopedia of plant anatomy. Bd. 10.
Teil 1 (pp. 1–675). G. Borntraeger.
Sennikov, A., Khassanov, F., Ortikov, E.,
Kurbonaliyeva, M., & Tojibaev, K.S. (2023). The
genus Iris L. s.l. (Iridaceae) in the mountains of
Central Asia biodiversity hotspot. Plant Diversity of
Central Asia, 2(1), 1–104. http://doi.org/10.54981/
PDCA/vol2_iss1/a1
Skrypec, C., & Odintsova, А. (2014). Anatomical
structure of pericarp in Gladiolus imbricatus L.
and Iris sibirica L. (Iridaceae Juss.). Modern
Phytomorphology, 6, 257–258. (In Ukrainian)
Skrypec, C., & Odintsova, А. (2015). Fruit and seed
morphology in Iris sibirica L. and Gladiolus imbricаtus L.
in relation with the modes of dissemination.
Biological Systems, 7(1), 93−96. (In Ukrainian)
Skrypec, K., & Odintsova, А. (2020). Morphogenesis
of fruits in Gladiolus imbricatus and Iris sibirica
(Iridaceae). Ukrainian Botanical Journal, 77(3), 210–
224. https://doi.org/10.15407/ukrbotj77.03.210.
(In Ukrainian)
Stoneburner, A.L., Meiman, P.J., Ocheltree, T.W.,
Nissen, S.J., & Bradfield, S.J. (2021). Simulated
trampling by cattle negatively impacts invasive
yellow-flag iris (Iris pseudacorus) when submerged.
Invasive Plant Science and Management, 14(4), 232–
239. https://doi.org/10.1017/inp.2021.28
Thadeo, M., Hampilos, K.E., & Stevenson, D.W.
(2015). Anatomy of fleshy fruits in the monocots.
American Journal of Botany, 102(11), 1–23. https://
doi.org/10.3732/ajb.1500204
Van Tieghem, M.P. (1871). Recherches sur la structure
du pistil et sur l’anatomie comparée de la fleur.
Mémoires présentés par divers savants à l’Académie des
sciences de l’Institut impérial de France, Séries 2, 21, 1–261.
Wilson, C.A. (2006). Patterns in evolution in
characters that define Iris subgenera and
sections. Aliso: A Journal of Systematic and
Evolutionary Botany, 22(1), 425–433. https://doi.
org/10.5642/aliso.20062201.34
Wilson, C.A. (2009). Phylogenetic relationships
among the recognized series in Iris section
Limniris. Systematic Botany, 34(2), 277–284. https://
doi.org/10.1600/036364409788606316
Zhygalova, S.L. (2014). Yellow iris (Iris
pseudacorus L.) in the flora of Ukraine: chorology.
In: V.V. Konischuk (Ed.), Ecology of wetlands
and peatlands (collection of scientific articles)
(pp. 93–95). Interservis. (In Ukrainian)
Мікроморфологія та анатомія плоду Iris pseudacorus L.
Анастасія Одінцова *, Ярослав Хомей
Львівський національний університет імені Івана Франка, вул. Грушевського, 4, Львів 79005, Україна;
* anastasiya.odintsova@lnu.edu.ua
Ґрунтуючись на уявленнях про те, що квітка і плід є цілісною еволюціонуючою структурою, ми
спробували з’ясувати особливості будови квітки, які проявляються у структурі плоду на прикладі
Iris pseudacorus, широко поширеного в Україні прибережного виду. Ми мали намір порівняти
результати з іншими видами Iris, дослідженими раніше, і виявити стабільність мікроморфологічних
ознак внутрішньої будови плоду. Для цього дослідження ми вивчали екзоморфологічні та
мікроморфологічні особливості зав’язі плоду, використовуючи модель вертикальної зональності
https://doi.org/10.15407/ukrbotj79.03.169
https://doi.org/10.15407/ukrbotj79.03.169
https://doi.org/10.3791/51381
https://doi.org/10.3791/51381
http://doi.org/10.54981/PDCA/vol2_iss1/a1
http://doi.org/10.54981/PDCA/vol2_iss1/a1
https://doi.org/10.15407/ukrbotj77.03.210
https://doi.org/10.1017/inp.2021.28
https://doi.org/10.3732/ajb.1500204
https://doi.org/10.3732/ajb.1500204
https://doi.org/10.5642/aliso.20062201.34
https://doi.org/10.5642/aliso.20062201.34
https://doi.org/10.1600/036364409788606316
https://doi.org/10.1600/036364409788606316
Plant Introduction • 99/100 23
Micromorphology and anatomy of fruit in Iris pseudacorus L.
гінецею, васкулярну анатомію та анатомію оплодня. У зав’язі I. pseudacorus на стадії плоду ми виявили
три вертикальні зони: коротку синасцидіатну, довгу симплікатну зону з насінинами, розміщеними
однорядно, та гемісимплікатну зону у носику плоду. Провідна система зав’язі складається з
дорзальних, септальних і вентральних жилок. Кожна з трьох дорзальних жилок поділяється
радіально на слід зовнішнього листочка оцвітини, слід тичинки і дорзальний пучок плодолистка.
Кожна септальна жилка поділяється тангентально на три пучки сліду внутрішнього листочка
оцвітини. Парні вентральні жилки входять у зав’язь знизу і живлять насінні зачатки і насінини.
Екзокарпій складається з багатокутних клітин з потовщеними целюлозними стінками. Ендокарпій
однорядний, нездерев’янілий, складається з прозенхімних клітин, витягнутих горизонтально.
В паренхімному мезокарпії виявлено велику кількість секреторних канальців з таніноподібним
вмістом. Розкривання плоду на три стулки здійснюється завдяки дорзальним та вентральним
щілинам. Дорзальні щілини формуються вздовж дорзальних борозенок і з’являються завдяки
дрібно-клітинній тканині навколо дорзальної жилки. Наявність вентральних швів плодолистків у
симплікатній зоні зав’язі забезпечує вентральне розкривання плоду. Структура плоду I. pseudacorus
подібна на структуру плодів інших видів роду Iris. Наше дослідження підтверджує, що вертикальну
зональність гінецею, плацентацію та провідну систему гінецею в роді Iris можна належним чином
проаналізувати на стадії плодоношення, оскільки компоненти будови зав’язі, які формуються на
пре-антетичній стадії, зберігаються у плоді.
Ключові слова: Iris, васкулярна анатомія, симплікатна зона, оплодень, зав’язь плоду
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| id | oai:ojs2.plantintroduction.org:article-1628 |
| institution | Plant Introduction |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2025-07-17T12:54:16Z |
| publishDate | 2023 |
| publisher | M.M. Gryshko National Botanical Garden of the NAS of Ukraine |
| record_format | ojs |
| resource_txt_mv | wwwplantintroductionorg/56/23058ac64c5754b647f18deb43598b56.pdf |
| spelling | oai:ojs2.plantintroduction.org:article-16282024-04-07T19:57:15Z Micromorphology and anatomy of fruit in Iris pseudacorus L. Мікроморфологія та анатомія плоду Iris pseudacorus L. Odintsova, Anastasiya Khomei, Yaroslav Based on the suggestion that flower and fruit are integrally evolving structures, we aimed to reveal the floral traits persisting in the fruit structure in Iris pseudacorus, a widely distributed riparian species in Ukraine. We intended to compare the results with the other Iris species studied previously and reveal the constancy of micromorphological features of fruit interior structure. We revealed exomorphological and micromorphological peculiarities of the fruiting ovary using the model of vertical zonality of the gynoecium, vascular anatomy, and general anatomy of the fruit wall. In the fruiting ovary of I. pseudacorus, we revealed the presence of three vertical zones: short synascidiate zone, long symplicate zone bearing uniseriate seeds, and hemisymplicate zone located in the fruit beak. The vascular system of the ovary is composed of dorsal, septal, and ventral veins. Each of three dorsal veins divides radially into the outer tepal trace, stamen trace, and dorsal carpellary bundle, while each septal vein divides tangentially into three bundles of the inner tepal trace. Paired ventral veins enter the ovary from its bottom and supply ovules and seeds. The exocarp is composed of polygonal cells with thickened cellulose walls. The endocarp is uniseriate, unlignified, and composed of live prosenchymal cells, which are elongated tangentially. In the parenchymatous mesocarp, a great number of secretory canals with tannin-like content occur. The dehiscence of fruit on three valves proceeded by both dorsal and ventral slits. Dorsal slits are formed along dorsal grooves and provided by small-celled tissue surrounding the dorsal veins. The presence of ventral sutures of carpels in the symplicate zone of the ovary provides ventral dehiscence of fruit. Hence, the structure of the fruiting ovary in I. pseudacorus is comparable to that of other Iris species. Our investigation confirmed that the vertical zonality, placentation, and vascular system of the gynoecium in Iris can be appropriately estimated in the fruiting stage because the structural components of the ovary, which developed at the pre-anthetic phase, persist in the fruit. Ґрунтуючись на уявленнях про те, що квітка і плід є цілісною еволюціонуючою структурою, ми спробували з’ясувати особливості будови квітки, які проявляються у структурі плоду на прикладі Iris pseudacorus, широко поширеного в Україні прибережного виду. Ми мали намір порівняти результати з іншими видами Iris, дослідженими раніше, і виявити стабільність мікроморфологічних ознак внутрішньої будови плоду. Для цього дослідження ми вивчали екзоморфологічні та мікроморфологічні особливості зав’язі плоду, використовуючи модель вертикальної зональності гінецею, васкулярну анатомію та анатомію оплодня. У зав’язі I. pseudacorus на стадії плоду ми виявили три вертикальні зони: коротку синасцидіатну, довгу симплікатну зону з насінинами, розміщеними однорядно, та гемісимплікатну зону у носику плоду. Провідна система зав’язі складається з дорзальних, септальних і вентральних жилок. Кожна з трьох дорзальних жилок поділяється радіально на слід зовнішнього листочка оцвітини, слід тичинки і дорзальний пучок плодолистка. Кожна септальна жилка поділяється тангентально на три пучки сліду внутрішнього листочка оцвітини. Парні вентральні жилки входять у зав’язь знизу і живлять насінні зачатки і насінини. Екзокарпій складається з багатокутних клітин з потовщеними целюлозними стінками. Ендокарпій однорядний, нездерев’янілий, складається з прозенхімних клітин, витягнутих горизонтально. В паренхімному мезокарпії виявлено велику кількість секреторних канальців з таніноподібним вмістом. Розкривання плоду на три стулки здійснюється завдяки дорзальним та вентральним щілинам. Дорзальні щілини формуються вздовж дорзальних борозенок і з’являються завдяки дрібно-клітинній тканині навколо дорзальної жилки. Наявність вентральних швів плодолистків у симплікатній зоні зав’язі забезпечує вентральне розкривання плоду. Структура плоду I. pseudacorus подібна на структуру плодів інших видів роду Iris. Наше дослідження підтверджує, що вертикальну зональність гінецею, плацентацію та провідну систему гінецею в роді Iris можна належним чином проаналізувати на стадії плодоношення, оскільки компоненти будови зав’язі, які формуються на пре-антетичній стадії, зберігаються у плоді. M.M. Gryshko National Botanical Garden of the NAS of Ukraine 2023-09-24 Article Article application/pdf https://www.plantintroduction.org/index.php/pi/article/view/1628 10.46341/PI2023004 Plant Introduction; No 99/100 (2023); 12-23 Інтродукція Рослин; № 99/100 (2023); 12-23 2663-290X 1605-6574 10.46341/PI99-100 en https://www.plantintroduction.org/index.php/pi/article/view/1628/1544 Copyright (c) 2023 Anastasiya Odintsova, Yaroslav Khomei http://creativecommons.org/licenses/by/4.0 |
| spellingShingle | Odintsova, Anastasiya Khomei, Yaroslav Мікроморфологія та анатомія плоду Iris pseudacorus L. |
| title | Мікроморфологія та анатомія плоду Iris pseudacorus L. |
| title_alt | Micromorphology and anatomy of fruit in Iris pseudacorus L. |
| title_full | Мікроморфологія та анатомія плоду Iris pseudacorus L. |
| title_fullStr | Мікроморфологія та анатомія плоду Iris pseudacorus L. |
| title_full_unstemmed | Мікроморфологія та анатомія плоду Iris pseudacorus L. |
| title_short | Мікроморфологія та анатомія плоду Iris pseudacorus L. |
| title_sort | мікроморфологія та анатомія плоду iris pseudacorus l. |
| url | https://www.plantintroduction.org/index.php/pi/article/view/1628 |
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