HISTO-ANATOMICAL OBSERVATIONS REGARDING …...Anca 1MEREACRE *, Angela TONIUC1, Constantin TOMA1...
Transcript of HISTO-ANATOMICAL OBSERVATIONS REGARDING …...Anca 1MEREACRE *, Angela TONIUC1, Constantin TOMA1...
Analele Ştiinţifice ale Universităţii „Al. I. Cuza” Iaşi
s. II a. Biologie vegetală, 2014, 60, 1: 13-24
http://www.bio.uaic.ro/publicatii/anale_vegetala/anale_veg_index.html
ISSN: 1223-6578, E-ISSN: 2247-2711
HISTO-ANATOMICAL OBSERVATIONS REGARDING VIOLA L. SPECIES IN
THE GÂRBOAVELE RESERVE (COUNTY OF GALAŢI)
Anca MEREACRE1*, Angela TONIUC1, Constantin TOMA1
Abstract: The authors research the structure of vegetative organs of five different vernal species of Viola
L. which grow in the Gârboavele - Galaţi nature reserve. The histological features, most of them
quantitative ones, which differentiate the five species of Viola L., are pointed out: the thickness of the
root and rhizomes peridermis, the number of palisadic layers of the leaf lamina, the presence or absence
of tector hairs and of oxaliferous cells, their frequency and position.
Key words: Viola, histo-anatomy, vegetative organs, Gârboavele-Galaţi reserve.
Introduction
There are 28 species of Viola L. in the Romanian flora (Grințescu et al., 1955; Sârbu
et al., 2013), four of which known to grow in the Gârboavele - Galaţi nature reserve: Viola
arvensis Murray, V. elatior Fr., V. hirta L. and V. odorata L. (Mititelu et al., 1968); to the
above mentioned ones we bring a 5-th one, recently found by us: V. kitaibeliana Schult.
Two of the species are annual ones (V. arvensis and V. kitaibeliana), while three of
them are perennial plants (V. elatior, V. hirta and V. odorata), and have a thick rhizome,
with or without aerial runners (V. hirta).
The anatomic structure of Viola species has been little investigated until now, as can
be seen from the two synthesis treaties, the one about the anatomy of dicotyledonous plants
(Metcalfe and Chalk, 1972) and the one about the leaves of angiosperms in general (Napp-
Zinn, 1973, 1974). When presentations of tissue structures of various vegetative organs are
made, the authors mention, among others the Viola genre, too; Viola odorata being most
frequently quoted.
Burduja and Moţiu (1982) achieve a detailed histo-anatomical study of the aerial
parts of Viola hymettia Boiss., a species closely related (often mistaken for ) to V. arvensis,
V. tricolor and V. kitaibeliana.
The aerial parts of V. arvensis are used (besides V. tricolor) in traditional medicine
for their anti-inflammatory, expectorant and diuretic and other properties. The need to
know more about the morpho-anatomic features of both species resulted in research work
dedicated to these aspects (Meyer, 1916; Toiu et al., 2009, 2010).
Reference to structure features of Viola species can be found in works of
comparative (Petit, 1887; Howard, 1962; Uphof and Hummel, 1962) or ecological anatomy
(Constantin, 1885), but also in some with a special focus on vernal species (Keller, 1934;
Ubaidulaev, 1959; Ivanskaja, 1962, 1963; Goryşina, 1965).
1 Faculty of Biology, “Al. I. Cuza” University, Bd. Carol I, no.11, Iaşi – 700506 Romania;
[email protected] (corresponding author*)
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Mention must be made that three of the species studied by us (Viola arvensis, V.
kitaibeliana and V. odorata) have been analyzed from an anatomical point of view besides
other plants growing in Northern Iran (Yousefi et al., 2012).
From the above we can conclude that there are few histo-anatomical data referring to
Viola L. species, reason why, in our work, we are attempting a comparative analysis of the
vegetative organs of the five vernal Viola species mentioned above, which grow in the
Gârboavele - Galaţi nature reserve.
Materials and methods
The material under study was collected in the Gârboavele - Galaţi nature reserve in
May 2012, at the anthesis stage. This forest-park is at 90 m altitude, and are predominantly:
Quercus pubescens, Q. pedunculiflora, Ulmus foliacea, Tilia tomentosa, to which are added
Acer campestre, A. tataricum, Pyrus pyraster, Cotinus coggygria and another species
(Mititelu et al., 1968).
The plants were fixed and preserved in 70% ethyl alcohol. The vegetative organs
(the root, rhizome, aerial runner, aerial stem, and leaf) were sectioned and coloured (in
iodine green and ruthenium red) according to the usual histo-anatomical technique in plant
research. Cross-sections and the tangential/longitudinal ones have been analyzed under the
Olympus CX31 optic microscope and photographed with an Olympus C 5060 camera
(Andrei and Predan, 2003).
Results and discussions
THE ROOT (Plate I, Figs. 1-3)
The rhizodermis of V. odorata and V. hirta occasionally presents long absorbent
hairs; the rhizodermis, as well as the external layer of the bark of the other three species is
exfoliates early.
The bark of V. odorata and V. hirta is differentiated in the exodermis, cortical
parenchyma and endodermis; the other species keep only the internal layers of the cortical
parenchyma (at V. hirta with aerial gaps) and the casparian type of endodermis; some of the
parenchymatous cells of V. hirta and V. odorata contain calcium oxalate ursines.
The central cylinder is the anatomical area with a secondary structure, due to the
activity of the vascular cambium which has produced a thin external phloem ring and a
thick central xylem body. The phloem ring consists of sieve-tubes, companiong cells and
numerous cells of amiliferous parenchyma (collenchymatous in V. arvensis, arranged in
strictly radial rows in V. elatior). Only Viola odorata has an incipient secondary structure,
the phloem has few parenchyma cells, and the xylem has very few libriform elements.
The central xylem body comprises vessels of varied diameters, irregularly dispersed
in the fundamental lignified mass. The axial part of the stem of V. arvensis, V. hirta and V.
kitaibeliana is less lignited, wih fewer and smaller vessels which are separated only by
cellulosic parenchyma.
The species were determined by Ion Sârbu, Ph.D, whom we warmly thank in this paper, too.
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Thus, the early passage from the primary to the secondary structure can be noticed
both in perennial and in annual species, but only at the level of the central cylinder, on
account of the vascular cambium.
THE RHIZOME (Plate I, Fig. 5)
Only Viola hirta and V. odorata present rhizomes, both are perennial species, but
only one of them has aerial runners with adventitious roots from the nodes (V. odorata).
The contour of its cross section is very irregular, with ribs of various size in V. hirta
and a symmetrical structure in V. odorata. The structure of both species is typically
secondary, resulting from the activity of both lateral meristems, but especially of the
vascular cambium.
The epidermis and the largest part of the cortical parenchyma have exfoliated. In the
deep persistent internal parenchymatic layers of V. odorata some cells contain, like the ones
in the pith, calcium oxalate ursines.
The phellogen (or cork cambium), differentiated at the level of the internal cortical
parenchyma or of the pericycle (V. hirta), has produced one or two areas of suber and the
phellem. The vascular cambium has produced the secondary conducting tissues, of a
fascicular type in V. odorata and ring type in V. hirta.
The secondary phloem ring is relatively thick and contains conducting elements
(sieve-tubes, companion cells) in the close vicinity of the cambium and numerous cells of
amiliferous parenchyma towards the exterior, disposed in radial rows, some containing
calcium oxalate ursines.
The secondary xylem ring is much thicker, strongly sclerified and intensely
lignified, made up of irregularly dispersed vessels of varied diameters, many libriform
fibers and few parenchyma cells. The xylem ring is crossed by some parenchymatic-
cellulosic medullar rays
In the thickness of the V. hirta ring we can distinguish several (4-5) rings, each of
them having more numerous early vessels and libriform fibers with thinner walls, and fewer
later vessels and libriform fibers with thicker walls. In the early xylem areas we can see
some islands of cellulosic parenchyma. The pith is parenchymatic-cellulosic, and many
cells contain calcium oxalate ursines.
THE AERIAL RUNNER (Plate I, Fig. 4)
The aerial runner, which we find only in V. odorata has a flat-elliptical contour in
cross-section, with a slightly concave face (the inferior one), which corresponds to the place
where the endogenously formed adventitious roots will come out; here the phloem is
reduced and interrupted by the cord which forms the adventitious root.
When dealing with thinner and younger material, the contour of the cross-section is
circular, the cortical parenchyma is exfoliating, the suber zone has 3-4 layers of suberified
cells, the phellodermis area is visibly collenchymatized, and the phloem ring has a different
thickness on the circumference of the organ. The structure is typically secondary and
asymmetrical for thicker and older material.
The aerial runner surface is covered by a relatively thick peridermis, with 4-6 layers
of suber (exfoliating) and an equal number of phellodermis layers, which are tangentially
elongated and are disposed in radial series, some of them containing calcium oxalate
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ursines. From place to place there are remains of the primary cortical parenchyma, adhering
to the suber.
The central cylinder is very thick, but irregular thickness along the circumference of
the aerial runner, so that it gets the shape of a cordiform contour in cross-section, the xylem
getting into direct contact with the suber along one of the sides. We see that the cambium
produces an incomplete, but thick, phloem ring (sieve-tubes and companion cells) towards
the interior and numerous parenchyma cells, disposed in radial rows and with moderately
thickened walls, towards the exterior.
Cambium also produces a thick, completely lignified, xylem ring (vessels, libriform
fibers, parenchyma cells), pierced by 1-2 parenchymatic-cellulosic medullar rays. Many of
medullar rays cells, as well as those of the liberian parenchyma, contain calcium oxalate
ursines.
At a closer analysis one can notice, inside the secondary xylem ring, 6-7 incomplete,
concentric rings which can be recognized by the early vessels of a larger diameter and by
the terminal libriform areas.
THE AERIAL STEM (Plate II, Figs. 6-11)
The structure of the two annual species (V. arvensis and V. kitaibeliana) is and
remains only a primary one. The structure of the three perennial species (V. elatior, V. hirta
and V. odorata) is mostly of secondary origin in the lower third of the stem, but only at the
level of the central cylinder, due to the activity of the vascular cambium.
The contour of the cross-section varies from circular to elliptically-circular
(modified by two lateral wings (V. elatior), to trapezoid (V. arvensis) or square shaped at
the tip and triangle shaped at the basis of the stem (V. kitaibeliana).
The epidermis has isodiametric cells, with thicker internal and external walls than
the other ones; the external wall is covered by a thin, lightly striped cuticle. From place to
place, at least in the upper third of the stem, we can notice stomata and tector hairs. There
are numerous unicellular relatively short hairs on the lateral wings in the lower third of the
stem in V. arvensis, V. elatior and V. kitaibeliana.
The bark is parenchymatic, of meatic type, but it also has aerial gaps (V. elatior, V.
odorata), and it ends with an endodemoid (V. arvensis) or with a casparian endodermis (V.
hirta, V. odorata). The hypodermic layer is collenchymatized in the ribs most frequently of
an angular or tangential type (V. kitaibeliana).Cells with calcium oxalate ursines have been
notice in V. elatior, V. hirta and V. odorata.
The central cylinder of annual species has conducting tissue of fascicular type (with
a primary structure) while the perennial species have ring type tissue (towards the basis of
the stem, with a secondary structure). The number of vascular bundles varies from the tip
(4) towards the basis of the stem (8-12) in V. kitaibeliana, it is constant (4) in V. arvensis or
it is higher (8-10) at the tip of the stem. In perennial species the vascular bundles unite
giving birth to two concentric rings at the stem basis.
In the primary structure the phloem has sieve-tubes and companion cells, while the
xylem has vessels and cellulosic parenchyma type of cells. The secondary phloem ring has,
in addition to this, parenchyma cells, and the secondary xylem ring has also libriform
fibers. The medullar rays, sclerified and lignified, form, together with the secondary xylem,
Mereacre, A. et al. 2014/ Analele Stiint. Univ. Al. I. Cuza Iasi, Sect. II a. Biol. veget., 60, 1: 13-24
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a thick and compact ring. We can observe thin sclerenchymatic fiber cords on the external
face of the phloemic ring (V. elatior).
The pith is parenchymatic-cellulosic and many of its cells contain calcium oxalate
ursines (V. odorata, V. elatior, and V. hirta). Many pith cells of the annual species become
disorganized thus creating a smaller or larger central aerial cavity, along the whole stem.
Such small aerial cavities in the pith have been noticed in V. elatior, too.
THE LEAF (Plates III, IV, V)
The petiole (Plate III, Figs. 12-17)
The contour of the cross-section varies from triangular (V. kitaibeliana) to
semicircular modified by two lateral-adaxial wings (three in V. odorata).
The epidermis has isodiametric cells, whose exterior walls are slight thicker than the
others and they are covered by a very thin cuticle. From place to place one can notice
stomata and rare unicellular tector hairs (more frequent on the lateral-adaxial wings),
shorter (V. elatior) or longer ones (V. hirta).
The fundamental parenchyma is slightly collenchymatised under the epidermis
(especially in the lateral-adaxial wings), with aerial cavities on adaxial face (V. arvensis, and
V. odorata); some of the cells contain calcium oxalate ursines (V. elatior and V. odorata).
The conducting tissues form 3 vascular bundles: a large median one and two small
lateral-adaxial ones, all of them with a primary structure.
The lamina (Plate IV, Figs. 18-21; Plate V, Figs. 22-26)
The epidermis, in front side view, presents cells with irregular contours, with slight
waving lateral walls on the upper face of the lamina and strongly waving ones on its lower
face; only V. kitaibeliana presents lateral walls with high amplitude waves on both faces of
the limb.
Anisocytic type stomata (cruciferous ones) can be noticed in the two types of
epidermis, with a higher frequency on the surface unit of the lower epidermis, this
indicating an amphystomatic leaf. In V. hirta the stomata are not present or are extremely
rare in the upper epidermis, indicating a hypostomatic leaf.
The mesophyll comprises two types of assimilating tissues: palisadic and spongy
ones, that the limb has a bifacial heterofacial structure (dorsiventral leaf) in all five Viola
species we have investigated. The differences between species refer to the number of
palisadic layers (one in V. hirta and two in the other species), to the length of the
composing cells (longer in V. kitaibeliana and shorter in V. elatior and V. hirta ), to the
presence or absence (V. arvensis ) of unicellular tector hairs, to the presence (V. hirta and
V. odorata) or absence of oxaliferous cells.
Conclusions
The general structure plan of all investigated vegetative organs is similar in all five
Viola species.
The passage from the primary to the secondary structure of the root takes place very
early both in perennial and annual species, but only at the level of the central cylinder, that
is on account of the vascular cambium.
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The rhizome, present only in perennial species (V. hirta and V. odorata) has a
secondary structure, as a result of the activity of both lateral meristems; the vascular
cambium and the cork cambium (phellogen). In the very thick secondary xylem ring we can
distinguish several areas, each with early and late wooden vessels (V. hirta). Both species
present many oxaliferous cells in the primary and medullar cortical parenchyma.
The aerial runner, present only in V. odorata, has, like the rhizome, a secondary
structure, with a thick peridermis, very thick xylem ring, oxaliferous cells in the medullar
rays, phelloderm and liberian parenchyma.
The five species we have investigated are differentiated only by the presence or
absence of tector hairs, of oxaliferous cells and central aerial cavity.
The leaf petiole has a different cross-section contour, with two lateral-adaxial wings
always visible. The species under study differ by the presence or absence of tector hairs,
oxaliferous cells and aerial cavities in the fundamental axial parenchyma. All five species
present 3 vascular bundles.
The foliar lamina has a homogenous, bifacial, heterofacial (dorsiventral) structure,
the difference between the species being mostly quantitative ones (the number of palisadic
layers, the length of their cells), more rarely qualitative ones: the presence or absence of
tector hairs and oxaliferous cells.
REFERENCES
Andrei, M., Predan, G.M.I., 2003. Practicum de morfología şi anatomía plantelor. Edit. Ştiinţelor Agricole,
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hymettia Boiss. et Heldr. Culeg. de studii şi artic. de Biol., Grăd. Bot. Iaşi. 2: 304-312.
Costantin, J., 1885. Recherches sur l’influence qu’exerce le milieu sur la structure des racines. Ann. des Sci. Nat.,
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Grinţescu, G., Guşuleac, M., Nyárády, E., 1955. Viola L. In Flora R.P.R., 3. Edit. Acad. Rom., Bucureşti: 553-
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Howard, R.A., 1962. The vascular structure of the petiole as a taxonomic character. Proc. 15-th Internat. Hort.
Congress, Nice, 1958: 7-13.
Ivanskaja, E.N., 1962. Sur quelques particularités de la structure anatomique de la feuille de plantes en rosettes et
en coussinet, d’altitude. Dokl. AN SSSR. 146, 2: 467-470.
Ivanskaja E.N., 1963. O stroenie mezofila list’ev nekotoryh vysokogornyh ratenij ţentral’nogo Kavkaza. Zapiski
Ţentral. Kavkazkogo otd. Vsesoiuz-go botanici-go obscestva, Ordjonikidze. 1: 87-97.
Keller E.F., 1934. Osobennosti anatomiceskogo stroenia list’ev u vesennih efemero-odnoletnikov. Sovremennaia
botanika. 2, 4: 83-95.
Metcalfe, C.R., Chalk, L., 1972. Anatomy of the Dicotyledons, 1. Clarendon Press, Oxford: 102-108.
Meyer, F.J., 1914. Bau und Ontogenie des Wasserleitungssystems der vegetativen Organe von Viola tricolor var.
arvensis. Thesis, Marburg (cf. Bot. Zbl, 141, 1919: 211-212).
Mititelu, D., Gociu, Z., Patraşcu, A., Gheorghiu, V., 1968. Flora şi vegetaţia pădurii-parc Gîrboavele-Galaţi.
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Napp-Zinn, Kl., 1973, 1974. Anatomie des Blattes. II. Blattanatomie der Angiospermen. Handbuch der
Pflanzenanatomie, Bd. VIII, 2 A 1-2, Gebrüder Borntraeger, Berlin.
Petit, L., 1887. Le petiol des Dicotyledones au point de vue de l’anatomie comparée et de la taxinomie. These,
Bordeaux.
Sârbu, I., Ştefan, N., Oprea, A., 2013. Plante vasculare din România. Edit. VictorBVictor, Bucureşti: 452-460.
Toiu, A., Oniga, I., Tămaş, M., 2009 Cercetări morfologice şi anatomice asupra speciei Viola tricolor L.
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Toiu, A., Oniga, I., Tămaş, M., 2010. Morphological and anatomical researches on Viola arvensis Murray
(Violaceae). Farmacia. 58, 5: 654-659.
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Tian-Şan. Avtoreferat, Taşkent.
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EXPLANATION OF THE PLATES
PLATE I
Fig. 1.Viola arvensis - cross-section through the root (Oc.10 x Ob.4)
Fig. 2. Viola kitaibeliana - cross-section through the root (Oc.10 x Ob.4)
Fig. 3. Viola hirta - cross-section through the root – details (Oc.10 x Ob.20)
Fig. 4. Viola odorata - cross-section through the aerial runner – details (Oc.10 x Ob.20)
Fig. 5. Viola hirta - cross-section through the rhizome (Oc.10 x Ob.4)
PLATE II
Fig. 6. Viola elatior - cross-section through the upper level of the stem (Oc.10 x Ob.10)
Fig. 7.Viola elatior - cross-section through the upper level of the stem – details (Oc.10 x Ob.20)
Fig. 8. Viola kitaibeliana - cross-section through the middle level of the stem (Oc.10 x Ob.10)
Fig. 9. Viola hirta - cross-section through the middle level of the stem (Oc.10 x Ob.10)
Fig. 10. Viola arvensis - cross-section through the basal level of the stem (Oc.10 x Ob.40)
Fig. 11. Viola elatior - cross-section through the basal level of the stem (Oc.10 x Ob.20)
PLATE III
Fig. 12. Viola.kitaibeliana - cross-section through the petiole (Oc.10 x Ob.10)
Fig. 13. Viola elatior - cross-section through the petiole (Oc.10 x Ob.10)
Fig. 14. Viola arvensis - cross-section through the petiole (Oc.10 x Ob.10)
Fig. 15. Viola odorata - cross-section through the petiole (Oc.10 x Ob.10)
Fig. 16. Viola elatior - cross-section through the petiole – details (Oc.10 x Ob.20)
Fig. 17. Viola hirta - cross-section through the petiole – details (Oc.10 x Ob.20)
PLATE IV
Fig. 18. Viola arvensis –the lower and upper lamina epidermis, in front view (Oc.10 x Ob.40)
Fig. 19. Viola elatior - the lower and upper lamina epidermis, in front view (Oc.10 x Ob.40)
Fig. 20. Viola hirta - the lower and upper lamina epidermis, in front view (Oc.10 x Ob.40)
Fig. 21. Viola odorata - the lower and upper lamina epidermis, in front view (Oc.10 x Ob.40)
PLATE V
Fig. 22. Viola arvensis - cross-section through the lamina in the midrib part (Oc.10 x Ob.20)
Fig. 23. Viola odorata - cross-section through the lamina in the midrib part (Oc.10 x Ob.20)
Fig. 24. Viola elatior - cross-section through the lamina in the midrib part (Oc.10 x Ob.20)
Fig. 25. Viola kitaibeliana - cross-section through the lamina in the midrib part (Oc.10 x Ob.10)
Fig. 26. Viola .hirta - cross-section through the lamina between the veins - details (Oc.10 x Ob.40)
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PLATE I
Figure 1. Viola arvensis Figure 2. Viola kitaibeliana
Figure 3. Viola hirta
Figure 4. Viola odorata
Figure 5. Viola hirta
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Figure 6. Viola elatior
Figure 7. Viola elatior
Figure 8. Viola kitaibeliana Figure 9. Viola hirta
Figure 10. Viola arvensis
Figure 11. Viola elatior
PLATE II
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PLATE III
Figure 12. Viola kitaibeliana Figure 13. Viola elatior
Figure 14. Viola arvensis
Figure 15. Viola odorata
Figure 16. Viola elatior Figure 17. Viola hirta
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PLATE IV
Lower epidermis Upper epidermis
Figure 18. Viola arvensis
Figure 19. Viola elatior
Figure 20. Viola hirta
Figure 21. Viola odorata
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PLATE V
Figure 22. Viola arvensis
Figure 23. Viola odorata
Figure 24. Viola elatior Figure 25. Viola kitaibeliana
Figure 26. Viola hirta