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624 AGRICULTURAL BIOLOGY, (SEL'skokhozyaistvennaya BIOLOGIYA) ISSN 0131-6397 (Russian Ed. Print) SOME FEATURES of EMBRYOLOGY O

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624 AGRICULTURAL BIOLOGY, (SEL'skokhozyaistvennaya BIOLOGIYA) ISSN 0131-6397 (Russian Ed. Print) SOME FEATURES of EMBRYOLOGY O AGRICULTURAL BIOLOGY, ISSN 2412-0324 (English ed. Online) 2018, V. 53, ¹ 3, pp. 624-633 (SEL’SKOKHOZYAISTVENNAYA BIOLOGIYA) ISSN 0131-6397 (Russian ed. Print) ISSN 2313-4836 (Russian ed. Online) UDC 633.811:582.71:581.3 doi: 10.15389/agrobiology.2018.3.624eng doi: 10.15389/agrobiology.2018.3.624rus SOME FEATURES OF EMBRYOLOGY OF Rosa spinosissima L., R. canina L. AND R. ½ damascena Mill. INTACT AND VIRUS-INFECTED PLANTS S.V. SHEVCHENKO, T.N. KUZMINA Nikita Botanical Gardens — National Scientific Center RAS, Federal Agency for Scientific Organizations, 52, ul. Ni- kitskii spusk, pgt Nikita, Yalta, Republic of Crimea, 298648 Russia, e-mail [email protected] ( corres- ponding author), [email protected] ORCID: Shevchenko S.V. orcid.org/0000-0002-0534-5548 Kuzmina T.N. orcid.org/0000-0002-4218-6080 The authors declare no conflict of interests Acknowledgements: Supported financially by Russian Science Foundation (grant ¹ 14-50-00079) Received April 5, 2017 A b s t r a c t Due to high decorative and economically valuable traits, roses have long been used in gar- den and park construction, and as medicinal and aromatic raw materials. Selection carried out with roses for many years, has resulted in more than 30 thousand cultivars. To create new forms and vari- eties, it is necessary to in-depth know species biology, including developments of generative ele- ments. This paper shows the study of male and female generative structure formation in two wild species of Rosa genus, Rosa spinosissima L. and R. canina L., naturally grown on the north-eastern slope of Mount Chatyr-Dag, and in two R. ½ damascena Mill. cultivars, which may be involved in breeding as parental plants. As a result, we have determined the main embryological types of genera- tive sphere formation in these species. It has been shown that the anthers are four-nested with two thecas, the wall of microsporangium is formed centripetal. A formed wall of microsporangium con- sists of epidermis, endothecium, 2-3 middle layers and tapetum, which is a derivative of a secondary parietal tissue. A tetrad of microspores is formed simultaneously. Differentiation mitosis in microspores leads to the formation of 2 celled pollen grains. Mature pollen grains are 2 celled with 3 pores. Micro- sporangium wall of the mature anther is represented by the epidermis covered with cuticle, fibrous endothecium and remnants of a middle layer. Morphogenicity and viability of mature pollen of these species and varieties have been established (abnormal pollen grains can be up to 30 %). Pollen sizes were measured. The diameter of pollen grains varies from 19 to 30 microns. The influence of viral infection on the formation of pollen grains has been shown, which causes an increase in the number of pollen abnormalities, changes the pollen grain size and ultimately leads to pollen grain degenera- tion. Defective pollen grains can be up to 50 %. Gynoecium of rose consists of many free carpels, each contains 1-2 ovules. Rosa ovule is anatropous, bitegmal, crassinucellate. Multicellular hypostase with thick-walled cells is formed in chalasa area. Micropyle is formed by an internal integument, which in its development is ahead of the outer one. Archesporium is polycelled and differentiated in the subepidermal layer of ovule. As a result of meiosis, linear tetrads of megaspores are mostly formed, of which chalasal or epichalasal megaspores were functionally active to derive embryo sacs of Polygonum-type. Mature embryo sacs are 8 nucleate and 7 celled. Egg apparatus consists of egg and 2 synergids, antipods are represented by three cells. Majority of embryo sacs gradually are obliterated, and only one of them matures. The studied species of genus Rosa are entomophilous plants pollinated by insects. It has been concluded that in selection, while choosing parental forms for hybridization, attention should be paid not only to the viability of their generative structures, but also to plant viral infection, since the presence of pathogens has led to a significant decrease in reproductive ability. Keywords: Rosa sp., male and female generative structures, viral infection, reproductive ability. Due to the variety of forms, abundant and long flowering, the rose has long been used in garden and park construction. The first information about the cultivation of roses was found for 4 thousand years BC; later, more than 30 thou- sand cultivars with decorative [1-3] and valuable medicinal characteristics [4-7] were created. Nowadays, the range of roses resistant to environmental stress fac- tors, diseases, and pests is expanding [8-10]. Cytological and genetic investigations 624 have shown the possibility to obtain polyploid forms [11-13]; the in vitro culture method has been widely used for the cultivation and reproduction of valuable gen- otypes of roses in recent decades [14-17]. It is important to mention that different species of the Rosa genus [18-20] can be used as medicinal plants. The Rosa genus has a wide geographic range, is widely studied and actively used in hybridization [21-24]. The genus includes 10 sections, including Pimpinel- lifoliae and Caninae, the representatives of which, along with others, serve as the original forms of garden varieties [6, 25-28]. The active development of industry and agriculture, the increase in air and soil pollution often lead to abnormal development of the generative sphere of plants; therefore, when conducting breeding, careful selection of parental forms is necessary. In this paper, the authors have presented the results of the investigation of the processes of male and female gametophyte formation in two prospective initial forms used in roses selection, Rosa spinosissima L. and R. canina L., and two high-yielding varieties of essential-oil-bearing rose (R.½ damascena Mill.), Dzhalita and Festivalnaya. In addition, the quality of pollen among visually healthy and virus-infected plants was assessed. The aim was to identify the features of the development and construc- tion of generative structures for the representatives of the three species from the Rosa genus in normal and viral infection. Techniques. Collection of samples from Rosa spinosissima and R. canina for making permanent preparations and observation of blooming were carried out in Crimea in 2016 under natural vegetation of the studied species in different populations on the North-Eastern slope of the Chatyr-Dag mountain at the alti- tude of 750-868 m above the sea level. The biomaterial of Dzhalita and Festival- naya cultivars (R. ½ damascena) was taken in the collection of Nikitsky botanical garden. To make permanent cytoembryological preparations, buds at different stages of development were fixed in the Carnoy mixture containing 96% aqueous solution of ethyl alcohol, chloroform, and glacial acetic acid in the 6:3:1 ratio. The material was dehydrated with butyl alcohol and xylene and then enclosed in paraffin [34, 35]. Paraffin serial sections with the thickness of 10-12 microns were prepared with a rotary microtome MRTU (Russia). The preparations were stained with methyl green-pironin with additional staining with Alcian blue [38] and viewed (light microscopes Jenaval and AxioScope A. 1, Carl Zeiss Micros- copy GmbH, Germany). PowerShot A550 digital camera (Canon, Inc., Malay- sia) and AxioCamERc5s image analysis system (Carl Zeiss Microscopy GmbH, Germany) were used for documentation. Temporary preparations of average pollen samples colored by 1% aceto- orcein were used for cyto-morphological characterization of pollen grains of R. canina, R. spinosissima, and R. ½ damascena. Morphologically normal and defective grains were counted. The first group includes the grains with homoge- neous coloring, a clearly expressed core of the vegetative cell, and the generative cell. Pollen grains with visual signs of cytoplasm and nuclei destruction of vege- tative and generative cells were considered defective. Their percent was determined as a ratio of the number of pollen grains with abnomalities to the total number of the recorded pollen grains for every species or variety. The di- ameter of pollen grains was measured on permanent preparations of average pol- len samples [37] (AxioVision Rel.4.8 software, Carl Zeiss Microscopy GmbH, Germany). Statistically significant differences in the proportion of defective pollen grains from anthers of R. ½ damascena with visual signs of infection and asymptomatic ones were determined by φ-angle transformation of Fischer (Fφ) 625 if p < 0.05 [36]. Data processing, with descriptive statistics and comparison of samples on the diameter of plants pollen grains with and without viral infection, was carried out with Statistica 6.0 (StatSoft Inc., USA), using Student's t- criterion. The arithmetic mean values (M) and standard errors of means (±SEM) are presented. Results. Rosa spinosissima L. (syn. Rosa pimpinellifolia L.) is a shrub up to 2 m with prickly shoots. Buzzes are of different sizes, thin, straight, usually ex- tended at the base. Leaves are with 5-11 folioles and naked stipules. Flowers are single, on long pedicels, sepals are simple, up to 15 mm long, remaining with fruits. Corollas are white, up to 5 cm in diameter, petals are sinuate, hypanthia are globular, baculums form a large head of the stigma. Fruits are up to 15 mm long, spherical or flattened-spherical, ripened, of dark color (up to black). De- spite the rather extensive geographic range, the population size of R. spinosissima is reduced, so it was listed in the Red Books of some regions, such as Kursk Re- gion and the Republic of Khakassia [29, 30]. R. spinosissima is the ancestor of many varieties, in particular, the extensive group of Scott's roses, which are known since 1600 and, according to modern ideas, belong to the Hybrid Spi- nosissima class [31]. In the past, many authors examined R. spinosissima and R. pimpinellifolia as different species, but molecular studies of samples from natu- ral populations of Great Britain have shown that they are of one species [10].
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