CC BY
65
Conclusion
In this work, the possible role of the Na+/Ca 2+ exchanger in the vasorelaxantion produced by 1-O-BK, a diterpenoid alkaloid, in rat aortic rings was elucidated for the first time. The present results support the idea that the 1-O-BK may relax vascular smooth muscle by a mechanism related to a decrease in [Ca 2+]i not only by inhibiting Ca 2+-influx through VDCCs or ROCCs, but also by blocking Ca 2+-entry via Na+/Ca 2+ exchanger. The finding that 1-O-BK
exhibits significant potency to block Ca 2+-influx via Na+/Ca 2+ exchanger may be important under pathological conditions where the exchanger, operating in reverse mode, induced Ca 2+ overload and, hence, may exacerbate overall vasoconstriction. Taken together, these data suggest that 1-O-BK is a promising compound for further development of novel approaches in the treatment of conditions associated with vascular smooth muscle disorders, such as hypertension or ischemia.
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14. Horiguchi S., Watanabe J., Kato H., Baba S., Shinozaki T., Miura M., et al. Contribution of Na+/Ca 2+ exchanger to the regulation of myogenic tone in isolated rat small arteries//Acta Physiol Scand. - 2001. - 173: 167-173.
15. Takai N., Yamada A., Muraki K., Watanabe M., Imaizumi Y. KB-R7943 reveals possible involvement of Na+/Ca 2+ exchanger in elevation of intracellular Ca 2+ in rat carotid arterial myocytes//J Smooth Muscle Res. - 2004. - 40: 35-42.
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Tursinbaeva Gulbakhor Sultanovna, Pedagogical State University name Nizami, Uzbekistan, Tashkent
Butnik Antonina Anatolievna, Institute of Gene Pool of Plants and Animals, Academy of Sciences
E-mail: [email protected]
Particular structure of fruits and seeds ephemers in the arid zone of Central Asia
Abstract: We describe the morphology and structure of the integuments of fruits and seeds 23 species from 18 genera, 5 families of ephemers from the most common (dominant) in the Kyzylkum desert. The next adaptive features were allocated: sclerification, slimy structure, water-carring tissue which promote the conservation of species in arid conditions. That was a negative correlation between the complexity of the pericarp and seed-coat structure. Keywords: morphology, anatomy, adaptive sign, diaspore, Kyzylkum.
Introduction
Ephemere plants are an important component of the vegetation in Central Asia deserts. In Kyzylkum 250 species from 20 families can be attributed to the ephemeral plants. The species of the families Asteraceae Dumort. Brassicaceae Burnett., Boraginaceae Juss. are most common [1, 336]. Ephemere plants are forming of ecological and biological diversity and occupied at the different ecological niches. Biomorphic of ephemere has several unique biological features.
Ephemere plants have a short-lived vegetation (1.5-2 months), but his latent period in the form of various diaspores (compound fruits, seeds) is longer and dependent on the external environment. They concentrated a considerable variety of adaptive traits [2, 160; 3, 35-38; 4, 36; 5, 29-30]. Diaspore (fruits and seeds) are carriers of the plant genome. They focused adaptive strategy tax-on [6, 376]. The study of the fruit and seeds of a necessary part in the knowledge of the ontogeny and adaptive strategy of plants.
Material and methods
Material collected in the natural habitat types in the southwestern Kyzylkum. The morphology, methods of dissemination and the anatomical structure of species of ephemere plants from the families Asteraceae Juss, Brassicaceae Burrneft, Boraginaceae Juss, Fabaceae Lindl., Lamiaceae Lindl. was studied.
The type of diasporas, their morphological parameters, the anatomical structure of the integument (pericarp, seed-coat) was described and reflected in the figures and micro photos. The stiff fruit was in mixture Strasburger: alcohol — glycerol — water (1:1:1). The structure of the fruit sketched drawing apparatus RA-6 and
Table 1. - The size of the
photographed. Description of the structure carried out by the usual method carpological studies on sections prepared manually [2, 160].
Results and discussion
Ephemer fruits are of different types in accordance with belonging to a particular family. For the family. Asteraceae is characterized achene with a tuft (pappus), with a rostellum and without (fig. 1a-c), a silicle is characters of the family Brassicaceae: indehiscent one-seeded (species Isatis), dispermous (Lachnoloma lechmannii, Goldbachia laevigata) (fig. 1 e-f), two-four seeds (species of the genus Alyssum) (fig. 1 g). Fruits of ephemeral Fabaceae (Astragalus, Onobrichis) (fig. 1 j-k) are one or a few seeds bean. Polyspermos pod at the different forms (round like, pear like) charactered for family Caryophyllaceae (Arenaria serpyllifolia, Spergu-laria microsperma) (fig. 1 h-i). The cenobium of the species family Lamiaceae are schizocarp fractional pod consisting of 3-4 fruts (Lallemantia royleana, Ziziphora tenuior) (fig. 1 L-m). Achene-shaped fruts with various appendages: small pricle, trichomes, eilaiosome, tulf (pappus) are the most widespread.
The dry fruits indeniscent oligospermous streamlined with alar and other appendages prevailed in desert conditions.
Reduction of the size and weight of fruits and seeds was promoted dominance in the desert of different forms of anemohorous dissemination. The fruit size of 1-20 mm. and a weight of 0.2-3.0 g. are dominating, seeds — 0.1-10 mm. and 0.05-20 g. (Table 1). The following forms dissemination is extended: anemochorous (evonemohorous and geohorous), ballisto-anemohorous, baro-horous, antitelahorous [7, 301-311; 8, 675]. The combination of 2 or more methods of dissemination often is observing (Table 1). diaspora ephemera (mm.)
Family, species Fruit Seed
Length Width Length Width
Asteraceae
Epilasia hemilasia (Bunge) Clark. 7.0 ± 0.6 1.8 ± 0.9 - -
Garhadiolus paposus Boiss. & Buhse 4.0 ± 0.3 1.5 ± 0.8 - -
Heteracia szovitsii Fish. &Mey. 2.9 ± 0.2 1.9 ± 0.9 - -
2.9 ± 0.2 0.7 ± 0.06 - -
Microcephala lamellata (Bunge) Polad. 4.0 ± 0.35 0.6 ± 0.05 - -
Senecio subdentatus Ledeb. 3.4 ± 0.3 0.4 ± 0.03 - -
Pullicaria gnaphalodus (Vent.) Boiss. 2.2 ± 0.2 0.5 ± 0.04 - -
Brassicaceae
Alyssum dasycarpum (Steph.) CAM 3.1 ± 0.3 3.0 ± 0.25 1.2 ± 0.1 0.9 ± 0.07
A. turkestanicum Regel & Shmalh. 3.6 ± 0.3 3.7 ± 0.3 1.6 ± 0.1 1.3 ± 0.1
A. szovitsianum Fish. & Mey 4.0 ± 0.35 3.6 ± 0.27 1.9 ± 0.1 1.5 ± 0.1
Goldbachia laevigata (Bieb.) DC. 8.4 ± 0.4 4.4 ± 0.4 2.7 ± 0.1 1.7 ± 0.1
Hymenolobusprocumbens (L.) Fourr. 4.1 ± 0.2 2.1 ± 0.2 0.6 ± 0.02 0.4 ± 0.03
Isatis minima Bunge 10.3 ± 1.3 2.8 ± 0.3 2.6 ± 0.2 1.0 ± 0.01
I. viollascens Bunge 10.6 ± 1.6 5.5 ± 0.4 3.4 ± 0.3 1.5 ± 0.01
Lachnoloma lechannii Bunge 5,6 ± 0.5 4.0 ± 0.4 3.5 ± 0.3 2.1 ± 0.02
Fabaceae
Astragalus ammophilus Kar. et Kiz 8.0 ± 0.75 2.5 ± 0.2 2.0 ± 0.1 1.0 ± 0.01
A. harpilobus Kar. & Kiz 20.5 ± 2.1 2.5 ± 0.25 2.2 ± 0.2 2.0 ± 0.02
A. cinarescens M. Pop. 7.0 ± 0.65 3.0 ± 0.27 1.6 ± 0.1 1.2 ± 0.01
Onobrichis tavernifolia Stocks & Boiss 11.7 ± 1.2 11.7 ± 1.25
Lamiaceae
Ziziphora tenuior L. calyx 6.9 1.8 ± 0.1 - -
ereme 1.7 0.6 ± 0.05 - -
Lallemantia royleana Benth. calyx 6.5 2.0 ± 0.15 - -
ereme 1.6 - - -
Fig. 1. Morphological types of fruit ephemera: achene — a — Heteraciaszovitsii(outdoor and indoor); b — Microcephala lamellate; c — Senecio subdentatus; silicle: dry-seeded indehiscent box — d — Isatis violascens; e — bispermous — Lachnoloma lehmanii; f — Goldbachia laevigata, 2-4 seeded dehiscent g — Alyssum dasycarpium; polyspermous box:
h — Spergularia microsperma; i — Arenaria serpyllifolia, monospermous seed bean: J - Onobrychis tavernifolia; k — Astragalus ammophilus; cenobia: skhizokarpnaya fractional box. L — Ziziphora tenuior; m — Lallemantia royleana
Fig. 2. The structure of pericarp and seed-coat of the fruits on the transverse section: a — Heteracia shovitsii; b — Senecio subdentata; c — Allysum szovitsianum; d — Euclidium syriacum; e — Isatis minima; f — Lallemanthia royleana. Abbreviation: Pr — pericarp, Sc — seed-coat, Wc — water-carring cell
Family Asteraceae. Fruit Amberboa turatica is ribbed achenes without rostellum, but multiserial pappus and elaiosome in the root part. Pericarp consists of a thick-walled parenchyma cells. Seed-coat thicker of pericarp at 2 times: 1 layer from 1-3 row sclerenchyma, 2-layer — 5-7 row parenchyma. Water-carring cells with spiral pore are located in the radical part.
Fruit Epilasia hemilasia is strongly pubescent ribbed achene without roustellum, but with pappus and collous ring in trichomes bases. Pericarp is sclerophyllous between the ribs and with water-carring cells in the ribs. A seed-coat is thin, with 2-3 rows of parenchyma.
Fruit Yarphadiolus papossum is achene in a head. Two morphotypes of fruits are allocated, i. e. expressed heterocarpous.
Next signs characterize the pericarp outer achenes: small prickle on the epidermis, 2-5 order water-carring cell with spiral-netted, liber-cells layer of 4-5 and 10-12 rows of cells. Seed-coat is thin with any parenchymal layers.
Fruit Heteracia szovitsii is heteromorphic achene with pappus and without it. In pericarp developed water-carring tissue, in mesocarp — sclerenchyma. Seed-coat is thin, parenchymal, pigmented (fig. 2 a).
Fruit Microcephala lamellate is heteromorphic achene in a head, pubescent, slightly ribbed on the dorsal side. The epidermis of the pericarp pigmented, wall cells is thicken with slimy fingerlike excrescence. The water-carring cells with spiral thickening walls disposed in mesocarp. Seed-coat is thicker pericarp, pigmented, parenchymal (fig. 2 a).
Fruit Pulicaria gnaphalodes is achene with pappus, downy. Pericarp in the top is parenchymal, at the lower part (mesocarp) — scler-enchymatous with 4-6 row layers. Seed-coat is 2 parenchymal rows.
Fruit Senecio subdentatus is densely planted ribbed achene with a round disk on top, tufted and rostellum. Epicarp is parenchymal, mesocarp consists ofparenchyma between the ribs and the sclerenchyma in ribs. Seed-coat is thin and parenchymal (Fig. 2b).
Thus, achenes of Asteraceae species have such adaptive signs as pubescens, water-carring cells, sclerenchyma and parenchyma in the pericarp, simplified parenchymal seed-coat (except Amberboa turanica). Remains of the endosperm provides additional protection.
Family Boraginaceae. Fruit of the species p. Heliotropium (H. lasiocarpum and H. biannulatiforme) is dry cenobia — dehiscing regma. The ereme is tetramospermous located in recesses of carpo-basis. Pericarp of ereme is composed: exocarp with pubescences of simple trichomes; mesocarp with 2-3-row parenchyma; pigmented and sclerotized endocarp. Seed-coat is thin, parenhymatous.
Family Brassicaceae. Fruit of species Alyssum (A. dasycarpum, A. szovitsianum, A. turkestanicum) is rounded, wide-septale silicle, with fruit style. Silicle is 2-4 locular, dehiscent. The pericarp is sclerotized, but the seed-coat fulfils function of the protection and consisting of mucous parenchyma, epidermal layer, pigmental layer and the adj acent 2-layer endosperm (starchy and protein) (Fig. 2c).
Multiple-row seed-coat, including thicked sclerenchyma cells, protects the embryo Euclidium syriacum. This is complicating the germination of seeds (Fig. 2d).
Fruit Goldbachia laevigata is dry, indehiscent, bilocular, hearless silicle. Pericarp is composed of different arangement sclerenchyma fibers. The walls sclerenchyma cells are very thick and alternating with multilayer parenchyma. Seed-coat is 5-6 cell rows and from it 2-3 rows of tangential elongated parenchymal cells, pigment and aleurone layers.
Fruit Hymenolobus procumbens is angustiseptate dehiscent, multi-seeded silicle. Seed-coat is thin and consists of mucous and pigmentary layers.
Fruit Lachnoloma lehmanii is indehiscent dilocular 2 tomentose pubescent silicle. Pericarp is composed of epidermal cells with thickened walls, 4-5 parenchymal layers, sclerenchyma layers and inner too sclerotized epidermis. Seed-coat is 5 layers: wavilike epidermal cells, 2-3 rows of compressed parenchymal cells, pigment and al-euronue layers.
The fruit of the species Isatis (I. minima, I. viollascens) is indehiscent dry pubescent silicle with teroid outgrowths. Pericarp includes pigmental layer under the epidermis, water-carring cells with netted and round-chinked pores and sclerenchyma. Seed-coat is thin, consisting of 2-3 rows of parenchymal cells (Fig. 2e).
Fruit Tausheria lasiocarpa is nutlike indehiscent bilocular downy silicle. Pericarp is composed of the pigment epidermal layer with a thick outer wall; 3-4 rows of small cell pigmented parenchyma, mechanical (sclerenchyma) strands surrounded water-carring cells. Vascular bandes (2) are surrounded by sclerotized fibres. Seed-coat consists 5 layers: the epidermis, parenchymal cells, sclerenchymal and aleuronic layers.
Family Caryophyllaceae. The fruit of Spergularia sperguloides is a dry dehiscent pod in caryx, pubescenced of grandular trichomes. Pericarp is thin and chaff-like. Seed-coat is formed from external integument. Exotesta is thickened, the cavity of cells filled up at tannin. Inneral integument is reduction. Reserved tissue is perisperm, as other species of family [9, 59-74].
Family Fabaceae. Fruit is polyspermous (A. harpilobus) or olygospermous bean, dry, dehiscent or partially dehiscent. The structure of the pericarp of studied species Astragalus (A. ammophi-lus, A. harpilobus, A. cinerascens) is similar. It consists of a single-row exocarp downy various trichomes; 3-4 orders parenchymal cells of mesocarp with the vascular bundles, which surrounded water-car-ring cells. Endocarp consists of divergent arrangement sclerenchymal fibers (parchment layer). Seed-coat consists of 5 layers: palisade cells (malpigie-cells), hypodermis (osteosclereide or lageniform sclereid), parenchyma, aleuronic layer. The fruits species have general similarity and distinctive signs: thick layer of malpigie-cells (A. ammophilus), thick pigmental layer (A. harpilobus), and pubescent of pericarp with exerescence (emergence) (A. cinerascens).
Family Lamiaceae. Fruit Lallemantia royleana is fractional pod (schizocarp) in tirse. Calyx is ribbed, downy prickle hairs, parenchymal, with the vascular bundles in the ribs. The pericarp of ereme is composed of slimy cells of the epidermis, 2-3 rows of pigmented parenchyma (Fig. 2 f).
Fruit of Ziziphora tenuior is fractional pod, with densely pubescent calyx, ribbed. Trichomes are simple, pricly and glandular with 8-12 secretory cells. Vascular bundle surrounded by sclerenchyma, is in each rib. Pericarps compose mucous epidermis, 2 rows of sclerenchyma and 1-2 rows of pigmental parenchyma. Seed-coat is thin, claying.
Conclusion
The fruit as part of the body's systems are complex adaptive traits. N. Kaden [10, 496] believed, that "the fruit should be regarded as organ which has developed through adaptation to better protects the seed and the different ways of dissemination". No less important are those signs associated with the functions of germination [6, 376]. In their view, the analysis of adaptive traits fruits should focus on adaptation to the dissemination methods, the protective features of the adverse factors and structural specialization to germinate.
Those are adequate basic functions of fruit: to survive, to expansion and to give rise to offspring.
The one-small dominated seed among the fruit of species (88.8 %), dry, indehiscent fruit the size of 3-10 mm. Availability
specialized hydro-cells in the pericarp and seed-coat in Asteraceae evidence of convergence of this trait. An important feature of the adaptive trait is to water-carring cells, which may be part of the pericarp or seed-coat.
A. Alyavdina [11, 85-100], Alexanderov, Rozhanovskiy [13, 20], Savchenkov [12, 129-147], Korobkov [14, 1302-1325] noted a correlation between dehiscence fruit and the presence of slimy layer. The dehiscence fruits of the family Brassicaceae are often slimy layer. We undisclosed slime cells in undeniscne fruits inherent on the plants of dry habitats. When the morning dew is settling, slime-cells quickly covered with a hydrocolloid film, poorly permeable at water.
The slimy-cells in fruits are a relatively wide-spread phenomenon among ephemeres [12, 129-147; 15, 108-109; 16, 1100-1111]. Slimy-cells in the fruit and seeds characterized plants by experiencing water shortages. Each epidermal cell of fruits species Alyssum miksospermiya contains slimy substance. Sclerotized of covers is one of the important signs of ephemeres. The sclerotized structure correlates with long-continued (for several years) preservation of seeds germination.
The fruits of each family and genus have their complex distinctive and adaptive signs caused by their belonging to a particular taxon. However, the same living conditions contributed to the formation of convergent features.
One of the leading adaptive signs in the family Brassicaceae is slimy-cells (p. Alyssum. Hymenolobus), and water-carring cells
layer, combined with scleriphication. Slimy-cells of A. szovitsianum marked as colorless columns rising from the bottom wall and expanding fungiform. A. dasycarpum is colorless columns rising from the bottom walls of the cells with a small extension in the upper part.
Common adaptive features primitving apocarpous fruit species of Astragalus (Fabaceae) are imperfect type dehiscence. This allows to seeds a long stay in the pericarp. Bilocular, different types of tri-chomes and epidermal formations; sclerotized of pericarp and seed-coat are protect against environmental influences. The pubescent of exocarp (A. harpilobus, A. remanens) presents or nakes (A. ammo-philus). Mesocarp is parenchymal cells with scleriphfillous vascular bundles, accompanied water-carring cells with spiral point and pore (A. harpilobus, A. remanens). The endocarp is sclerotized with parallel and perpendicular to the direction of the fibers. Seed-coat consists of 5 layers of differing power as all Fabaceae. The endosperm is the form of unstructured film.
Other families in this study presented a large number (2-3) species. In the family Lamiaceae observed complete slime-cells and sclerotized pericarp, provided deep physiological dormancy of seeds.
In the family Boraginaceae presence a thick layer of the endosperm and significant sclerotized pericarp.
All these features provide good protection of the embryo, but embarrassing germination. Increased protection of embryo against adverse conditions, the formation varying complexity coat of is regarded as one of the directions of evolution [17, 344].
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