ABSTRACT:
In this study, the morphological and anatomical features were investigated of three taxon of Hippocrepis L. species which spreading naturally in Turkey. In the morphological part of the study, H. unisiliquosa subsp. unisiliquosa’s stem is erect or decumbent. The species H. ciliata’s Willd. stem is erect. H. multisiliquosa’s L. stem is decumbent. The fruit of H. multisiliquosa is much more convoluted than the other two taxa. There are cilia on the fruit of H. ciliata, but there are not found any cilia the other two taxa. The stem anatomy of all the studied taxa is hexagonal. In the stem cross section of H. ciliata 12-14 vascular bundle are found, H. unisiliquosa subsp. unisiliquosa 12-15 and H. multisiliquosa 12-13. Leaf vascular bundles are arranged regularly, H. unisiliquosa subsp. unisiliquosa have 18-20, H. ciliata have 6-8, H. multisiliquosa have 9-13 vascular bundles.
Keywords:
anatomy; Leguminosae; morphology
RESUMO:
Neste estudo, foram investigadas as características morfológicas e anatômicas de três táxons de espécies de Hippocrepis L. que se espalham naturalmente na Turquia. Na parte morfológica do estudo, o tronco de H. unisiliquosa subsp. unisiliquosa está ereto ou decumbente. A haste de H. ciliata é ereta. O tronco de H. multisiliquosa é decumbente. O fruto de H. multisiliquosa é muito mais enrolado do que o dos outros dois taxa. Há cílios sobre o fruto de H. ciliata, mas não foram observados cílios nos outros dois taxa. A anatomia do caule de todos os taxa estudados é hexagonal. Na seção transversal do tronco de H. ciliata, 12-14 feixes vasculares são encontrados; em H. unisiliquosa subsp. unisiliquosa, 12-15; e em H. multisiliquosa, 12-13. Os feixes vasculares foliares são dispostos regularmente, com H. unisiliquosa subsp. unisiliquosa apresentando 18-20 deles; H. ciliata, 6-8; e H. multisiliquosa, 9-13.
Palavras-chave:
anatomia; Leguminosae; morfologia
INTRODUCTION
Hippocrepis L. is the genus of Leguminosae family. The genus is distributed in various parts of the World. For example, there are 21 species in Spain, 6 species and 1 variety in Morocco, 5 species in Algeria and 1 species in Caucasia. However, in Turkey Hippocrepis is represented by 4 taxa belongs 3 species (Davis, 1984Davis PH. Flora of Turkey and the East Aegean Islands. Edinburgh: Edinburgh University Press; 1984. 9v.). There are some systematic, chromosomal and enzymatic studies about genus Hippocrepis (Fearn, 1972Fearn GM. The distribution of intraspecific chromosome races of Hippocrepis comosa L. and their phytogeographical significance, New Phytol. 1972;71:1221-5.; Lonn et al., 1995Lonn M, Prentice HC. The structure of allozyme and leaf shape variation in isolated, range-margin populations of the shrub Hippocrepis emerus (Leguminosae). Ecography. 1995;18:276-85.; Mansour et al., 1995Mansour A, Williams JM, Wainwright SJ, Hipkin CR. Nitroaliphatic compounds in Hippocrepis comosa L. and other legumes in the European flora. Phytochemistry, 1995;40:89-91.; Hipkin et al., 1999Hipkin CR, Salem MA, Simpson D, Wainwright SJ. 3-nitropropionic acid oxidase from horseshoe vetch (Hippocrepis comosa): a novel plant enzyme. Biochem J. 1999;340(Pt 2):491-5.; Rosselló et al., 2002Rosselló JA, Carmencebrián M, Mayol M, Testing taxonomic and biogeographical relationships in a narrow Mediterranean endemic complex (Hippocrepis balearica) using RAPD markers. Ann Bot. 2002;89(3):321-7.; Hennenberg and Bruelheide, 2003Hennenberg KJ, Bruelheide H. Ecological investigations on the northern distribution range of Hippocrepis comosa L. in Germany. Plant Ecol. 2003;166(2):167-88.).
In this paper, the anatomy and morphology of Hippocrepis unisiliquosa subsp. unisiliquosa (Forssk.) Bornm., Hippocrepis ciliata Willd. and Hippocrepis multisiliquosa L. was studied in order to understand the usefulness of these characteristics for systematic purposes. By examining these three taxa belonging to Hippocrepis, it is aimed to find out if there is another differentiating property of these taxa within each other.
MATERIALS AND METHODS
Plant specimens for anatomical studies were collected between 2011 and 2012 during the flowering period from natural populations (Table 1).
A total of 30 individuals obtained from subpopulations at different altitudes were used. For anatomical studies, plant specimens were fixed in 70% ethanol. The paraffin method (Algan, 1981Algan, G. 1981. Microtechnics for the Plant Tissues. Publ. Firat Univ. Science and Art Fac., Istanbul, 1:1-94.) was used for preparing cross-sections of root, stem and leaves of each sample. Roots of the same diameter have been used for cross sections. Transverse sections 15-20 µm were made using a sliding microtome. Slides stained with safranin-fast green. Slides were photographed with motorised Leica DM 300 microscope. The measurements of root, stem and leaf cell size of the species were taken using an ocular micrometer. Minimum, maximum, mean and standard deviation were determined.
RESULTS AND DISCUSSION
Morphological characteristics
All the investigated taxa are annual herbs with a tap-root. The stem of H. unisiliquosa subsp. bisiliqua is ascending to erect and clearly quadrangular. The stem is approximately 25-105 cm height, branched at and near the base, pubescent with curved hair. H. unisiliquosa subsp. unisiliquosa’s stem is erect or decumbent. While the stem is erect of H. ciliata species H. multisiliquosa’s stem is decumbent. H. unisiliquosa subsp. unisiliquosa’s stem is 17-26 cm, H. ciliata’s stem is 16-28 cm, H. multisiliquosa’s stem is 10-42 cm (Table 2). The fruits of the taxa are lomentoid and convoluted. The fruit of H. multisiliquosa is much more convoluted than the other two taxa. There are cilla on the fruit of H. ciliata, but there are not found any cillia the other two taxa. H. unisiliquosa subsp. unisiliquosa’s fruit has 4-8 seed H. ciliata’s fruit has 5-8 seed, H. multisiliquosa’s fruit has 7-12 seed (Figures 1 and 2).
(A), (B): H. unisiliquosa subsp. unisiliquosa, (C): leaf of H. unisiliquosa subsp. unisiliquosa, (D): flower of H. unisiliquosa subsp. unisiliquosa, (E): fruits of H. unisiliquosa subsp. unisiliquosa, (F): seeds of H. unisiliquosa subsp. unisiliquosa (G), (H): H. ciliata, (I): leaf of H. ciliata, (J): flower of H. ciliata, (K): fruits of H. ciliata, (L): seeds of H. ciliata, (M), (N), (O): H. multisiliquosa.
Anatomical characteristics
When the root structure was examined in terms of its anatomical characteristics, a single row epidermis consisting of rectangular shaped cells was observed in all three taxa. The cortex layer in the root of H. unisiliquosa subsp. unisiliquosa is composed of 5-6 cell lines, in H. ciliata 9-13 cell lines, in H. multisiliquosa 10-12 cell lines. The stem anatomy of all the studied taxa is hexagonal. H. unisiliquosa subsp.. unisiliquosa and H. ciliata’s the length of epidermis cells are larger than their widths. In the other species is vice versa. The cortex layer of H. unisiliquosa subsp. unisiliquosa has 5-6, at H. ciliata 5-9, H. multisiliquosa 5-11 cell lines. In the stem cross section of H. ciliata 12-14 vascular bundle are found, H. unisiliquosa subsp. unisiliquosa 12-15 and H. multisiliquosa 12-13. The stem of all taxa investigated in this paper have large pith which are orbicular or oval parenchymatic cells with intercellular spaces and there is no fragmentation in the pith. When the leaf cross sections were examined, it was observed that all three taxa were of bifacial leaf type. The outer surface of the leaf blade is covered with a cuticle layer in all studied taxa. There is a single row of epidermis cells both upper and the lower face of all leaves. Lower epidermal cells are larger than upper epidermal cells in three taxa. The numbers of layers of palisade parenchyma cells are different in studied taxa, there are 1-2 layers in H. unisiliquosa subsp. unisiliquosa, there are 2-3 layers in the other two taxa. In taxon H. ciliata’s palisade parenchyma cells have intercellular space. In studied all taxa, the mesophyll is made up of elongated rectangular palisade parenchyma and isodiametric spongy parenchyma cells which contain very rich chloroplasts. The vascular bundles are arranged regularly, H. unisiliquosa subsp. unisiliquosa have 18-20, H. ciliata have 6-8, H. multi siliquosa have 9-13 vascular bundles. Spongy parenchyma cells are 1-2 layer in cross section of leaves of all studied taxa. H. unisiliquosa subsp. unisiliquosa and H. ciliata’s spongy parenchyma cells have larger intercellular spaces than H. multisiliquosa. All studied taxa are amphistomatic. When classified according to neighbouring epidermis cells, all types of stomata of the taxa are anomocytotic. According to friend cells, they have amaryllis type stomata (Figures 3, 4, and 5).
Anatomical cross sections of H. unisiliquosa subsp. unisiliquosa (A), (B): root, (C), (D), (E): stem, (F): leaf; c:cortex, e: epidermis, le: lower epidermis, p: pith, ph: phloem, pp: palisade parenchyma, sp: spongy parenchyma, ue: upper epidermis, x: xylem.
Anatomical cross sections of H. ciliata (A), (B): root, (C), (D): stem, (E), (F): leaf; c: cortex, e: epidermis, le: lower epidermis, p: pith, pp: palisade parenchyma, sp: spongy parenchyma, ue: upper epidermis, x: xylem.
Anatomical cross sections of H. multisiliquosa (A): root, (B), (C), (D): stem, (E), (F): leaf; c: cortex, e: epidermis, le: lower epidermis, p: pith, ue: upper epidermis, x: xylem.
In the current literature H. unisiliquosa subsp. unisiliquosa is referred to as H. unisiliquosa subsp. biflora. H. unisiliquosa subsp. bisiliquosa that is mentioned in Flora of Turkey was not found in any herbarium or field. The inability to find any permission for this subtype suggests the proposal of combining two subspecies. According to our findings morphological measurement of H. unisiliquosa subsp. unisiliquosa and H. unisiliquosa subsp. bisiliqua shows similarities. According to our findings the number of the flower of H. unisiliquosa subsp. unisiliquosa is 1-2, and their fruits have 4-8 seeds. According to Flora of Turkey the number of the flower of H. unisiliquosa subsp. bisiliqua 1-3 and their fruits have 6-10 seeds. There will be a definite decision after more detailed taxonomic examinations to be made (Table 3).
Tezcan (2008Tezcan S. Thermopsis turcica (Fabaceae) Kit Tan, Vural & Küçüködük Üzerinde Anatomik, Morfolojik ve Karyolojik Çalismalar Yüksek Lisans Tezi, Afyon Kocatepe Üniversitesi, Fen Bilimleri Enstitüsü; 2008.) studied on morphology, anatomy and karyology of Thermopsis turcica (Leguminosae). He found that rizoderma in roots of the species is composed of 6-8 rowed cells. Adjacent to the rizoderma cells, there are 13-19 layered cortex. They also said that there were pith rays from xylem tissue to phloem tissue and cambium rings are visible between xylem and phloem. It is reported that xylem occupied the very large area. The cortex layer at the root of H. unisiliquosa subsp. unisiliquosa consists of 5-6 cell lines, H. ciliata consists of 2-3 cell lines, H. multisiliquosa consist of the 10-12 cell lines. In the roots of all studied taxa, primer and secondary pith rays are observed. The very large area of the transverse section occurs with the component of xylem in all investigated taxa. It is stated that there is a regular sequence of epidermis in Thermopsis turcica’s stem. It has been reported that under the epidermis, there are single rowed collenchyma cells which are reaching up to ten rows at corners. And below them, there are 6-13 ordered parenchyma cells. It has been reported that there are phloem pith rays from cortex to xylem, cambium visible clearly, and the pith is composed of 7-10 rowed cells. The cortex region is composed of a 5-6 layer of cells in H. unisiliquosa subsp. unisiliquosa, 5-9 layer of cells in H. ciliata, a 5-11 layer of cells in H. multisiliquosa. It has been reported that at the transverse section of the leaf blade, upper and lower epidermis cells are different each other. There is an isolateral structure due to palisade parenchyma on the leaves, the sponge parenchyma cells have different shapes and because of the presence of stomata on both sides, leaf is amphistomatic and anomocytic stoma type was seen (Tezcan, 2008). In our study, it was observed that amphistomatic leaves and anomocytic stoma type in all three taxa.
REFERENCES
- Algan, G. 1981. Microtechnics for the Plant Tissues. Publ. Firat Univ. Science and Art Fac., Istanbul, 1:1-94.
- Davis PH. Flora of Turkey and the East Aegean Islands. Edinburgh: Edinburgh University Press; 1984. 9v.
- Fearn GM. The distribution of intraspecific chromosome races of Hippocrepis comosa L. and their phytogeographical significance, New Phytol. 1972;71:1221-5.
- Hennenberg KJ, Bruelheide H. Ecological investigations on the northern distribution range of Hippocrepis comosa L. in Germany. Plant Ecol. 2003;166(2):167-88.
- Hipkin CR, Salem MA, Simpson D, Wainwright SJ. 3-nitropropionic acid oxidase from horseshoe vetch (Hippocrepis comosa): a novel plant enzyme. Biochem J. 1999;340(Pt 2):491-5.
- Lonn M, Prentice HC. The structure of allozyme and leaf shape variation in isolated, range-margin populations of the shrub Hippocrepis emerus (Leguminosae). Ecography. 1995;18:276-85.
- Mansour A, Williams JM, Wainwright SJ, Hipkin CR. Nitroaliphatic compounds in Hippocrepis comosa L. and other legumes in the European flora. Phytochemistry, 1995;40:89-91.
- Rosselló JA, Carmencebrián M, Mayol M, Testing taxonomic and biogeographical relationships in a narrow Mediterranean endemic complex (Hippocrepis balearica) using RAPD markers. Ann Bot. 2002;89(3):321-7.
- Tezcan S. Thermopsis turcica (Fabaceae) Kit Tan, Vural & Küçüködük Üzerinde Anatomik, Morfolojik ve Karyolojik Çalismalar Yüksek Lisans Tezi, Afyon Kocatepe Üniversitesi, Fen Bilimleri Enstitüsü; 2008.
Publication Dates
-
Publication in this collection
19 Aug 2019 -
Date of issue
2019
History
-
Received
29 Aug 2017 -
Accepted
16 Oct 2017
