Steciana Doi:10.12657/Steciana.020.004 ISSN 1689-653X
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2016, Vol. 20(1): 21–32 Steciana doi:10.12657/steciana.020.004 www.up.poznan.pl/steciana ISSN 1689-653X ANATOMICAL STUDY OF CORNUS MAS L. AND CORNUS OFFICINALIS SEIB. ET ZUCC. (CORNACEAE) ENDOCARPS DURING THEIR DEVELOPMENT MARIA MOROZOWSKA, ILONA WYSAKOWSKA M. Morozowska, I. Wysakowska, Department of Botany, Poznań University of Life Sciences, Wojska Polskiego 71 C, 60-625 Poznań, Poland, e-mail: [email protected], [email protected] (Received: June 17, 2015. Accepted: October 20, 2015) ABSTRACT. Results of anatomical studies on the developing endocarps of Cornus mas and C. officinalis are pre- sented. Formation of an endocarp and anatomical changes in its structure from the flowering stage to fully developed fruits were observed with the use of LM and SEM. In the process of anatomical development of endocarps the formation of two layers, i.e. the inner and the outer endocarp, was observed. Changes in their anatomical structure consisted in a gradual thickening of cell walls and their lignification. The ligni- fication of endocarp cell walls begins in the inner endocarp, it proceeds in the outside parts of the outer endocarp, with an exception of several layers of cells forming the transition zone (circular strand) present on its margin, and finally, almost at the same time, in the rest of the outer endocarp. Cell walls of the cells distinguishing the germinating valves undergo thickening delayed by several days in comparison to other endocarp cells. Thickening of their cell walls starts in cells situated close to the inner endocarp and proceeds to its outer parts, and their lignification is not very intensive. The described scheme of the development and lignification of C. mas and C. officinalis endocarps is similar to that of Prunus cerasus (Rosaceae). KEY WORDS: dogwoods, fruit, anatomy, endocarp, lignification INTRODUCTION summer and autumn decorative bright-red fruits ma- The genus Cornus L. (Cornaceae) comprises approxi- ture. Cornus mas fruits thanks to the high contents of mately 58 tree and shrub species found mainly in the ascorbic acid and flavonoids are known for the high northern hemisphere in the temperate zone, rarely health-promoting value, particularly antioxidant ac- found in tropical Africa and South America (WANGE tivity (PAWLOWSKA et al. 2010). In Turkey they are used RIN 1910, MACBRIDE 1929, EYDE 1988, SENETA & DOLA for their antiallergenic, antihistaminic and bactericid- TOWSKI 2009, STEVENS 2012). al properties (ERCISLI et al. 2011). Vitamin C content In Poland only two wild species have been report- in fruit of this species is very high, over 100 mg/100 g ed, i.e. dwarf cornel (Cornus suecica L.), at present as- fruit, which is a value higher than that in other fruits signed the extinct species status (Żukowski & Jacko rich in that vitamin (VIDRIH et al. 2012). Moreover, its WIAK 1995), and common dogwood (Cornus sanguinea fruits also contain high levels of polyphenols and an- L.) (Bugała 2000). Many other species are grown and thocyanins (PANTELIDIS et al. 2007). Stones of corneli- planted as ornamentals in gardens and parks (MIREK an cherry fruits may be used as a raw material to pro- et al. 2002, Rutkowski 2006). These species include duce oil rich in unsaturated fatty acids (kuchaRska et e.g. Cornus mas L. (cornelian cherry) and Cornus of- al. 2009). Fruits of C. mas are also useful raw material, ficinalis Seib. et Zucc. (Asiatic dogwood or Japanese which may be processed to various products (DEMIR cornel dogwood). Decorative value of these species & kalyoncu 2003). In turn, fruits of C. officinalis are is connected with their early flowering, closely fol- used in traditional Chinese medicine to treat kidney lowing that of the common hazel and preceding for- disease and diabetic nephropathy. Studies have also sythia. At that time numerous tiny yellow-coloured shown a positive effect of glycosides isolated from flowers develop before leaves appear. Next in the late Cornus fruits against nephropathy. Loganin and its 22 Maria Morozowska, Ilona Wysakowska derivatives proved to be particularly effective in this MATERIAL AND METHODS respect (WEI MA et al. 2014). Both above-mentioned Cornus species investigat- Analyses were conducted on two species from the ed in this study belong to the group of ‘cornelian genus Cornus: C. officinalis and C. mas. Material for cherries’ (CC) comprising six species (Cornus sessilis investigations comprised fruits varying in their ma- Torr. ex Durand, C. mas L., C. officinalis Sieb. et Zucc, turity, which were collected from plants growing un- C. chinensis Wangerin, C. volkensii Harms, and C. ey- der comparable conditions at the Botanical Garden deana Q.Y. Xiang et Y.M. Shui). ‘Cornelian cherries’ of the Adam Mickiewicz University in Poznań and at constitute one of four major clades, into which Cor- the Arboretum in Kórnik, in western Poland (Table nus species are divided in the current phylogenetic 1). Plant material was sampled throughout the entire classification (XIANG et al. 1993, 1996, 1998, 2006, vegetation season in 2011, from the flowering phase FAN & XIANG 2001). to full fruit maturity, initially at 7-day and next at Detailed studies on the development and anatom- 14-day intervals. The successive stages in endocarp ical structure of the pericarp conducted to date con- development had to be analysed on different stones, cerned plants producing stone fruits from the fami- which had been growing under varied weather condi- ly Rosaceae (TUKEY & YOUNG 1939, KANIEWSKI 1963), tions, thus the total fruit formation time could only as well as selected species from the genus Rosa L. be roughly estimated. After harvest the experimen- (zieliński et al. 2010, GUZICKA et al. 2012). Moreover, tal material was preserved and stored in FAA (70% studies on the formation of the pericarp in several etha nol 90%, acetic acid 5%, formalin 5%). selected species from the genus Cornus were also car- In order to conduct an analysis of the anatomical ried out by KANIEWSKI & HAUSBRANDT (1968). In re- structure transverse sections (at three stone length lation to the genus Cornus a recent study described levels: at the base, at mid-section and under the and presented a comparative analysis along with an stone apex) and longitudinal sections were prepared analysis of genetic variation of the anatomical struc- both manually and using a Leitz sliding microtome. ture of mature fruits in C. mas and C. officinalis as well In order to detect lignified cell walls microtome slic- as their apparent hybrid (MOROZOWSKA et al. 2013). es of 50 μm in thickness were treated with floro- Results of the above mentioned studies indicate dif- glucinol with hydrochloric acid. Stained slides were ferent types of endocarp development in stone fruits. examined under a light microscope (LM) and images The thickening process followed by cell wall lignifi- of selected specimens were taken using an OptaView cation in the endocarp, despite certain regularities, camera. progresses differently in individual species. It varies Endocarp sections for observations under a scan- not only in terms of the site, at which it is initiated, ning electron microscope (SEM) were prepared but also the direction, in which it develops within manually, dehydrated in an acetone series at concen- the endocarp. trations of 30%, 50%, 70%, 90%, and 100%, three The aim of the presented study was to follow endo- times for 6 min each and sputtered with gold. Obser- carp development in C. mas and C. officinalis from the vations were conducted at the Laboratory of Electron moment of fruit setting to their full maturity, taking and Confocal Microscope, AMU in Poznań, under a into consideration the progressing process of thicken- Zeiss EVO 40 scanning electron microscope at 8–15 ing and lignification of cell walls in the fruit endocarp. kV depending on the species. Identification of individual stages in endocarp devel- Since fully developed stones of dogwood fruits are opment will show whether and if so, following which very hard, preparation of transverse sections turned of the patterns of drupe formation described to date, out to be impossible at their full maturity, i.e. after endocarps develop in the analysed species. reaching full colouring. Table 1. Location and provenance of analysed species Species Symbol of plant collection Plant number in plant collection Origin Cornus mas AMU BGP 191 no data AMU BGP 1519 planted before 1951 AMU BGP 7227 The Bolestraszyce Arboretum, Poland KA 49 planted before 1926 Cornus officinalis KA 6956 Kyoto Botanical Garden, Japan AMU BGP 04987 Dresden, Germany Explanation: AMU BGP – Adam Mickiewicz University Botanical Garden in Poznań, KA – The Kórnik Arboretum. Anatomical study of Cornus mas L. and Cornus officinalis Seib. et Zucc. (Cornaceae) endocarps... 23 RESULTS AND DISCUSSION changes were taking place within the stone, consist- ing in the increase in the thickness of cell walls and Analysed plants from both Cornus species differed their lignification, i.e. transformation into sclereids. slightly in the phenology of flowering and fruiting, In the above-mentioned period changes in fruit size, which may result from their genetic variation and lo- including their length, were not as marked. Through- cal weather conditions during the vegetation period. out July the process of fruit maturation was progress- However, the period of fruit formation from flow- ing, during which their colour was changing and the ering to the development of mature fruits and their fruits were becoming increasingly fleshy. In rela- complete colouring lasted for approximately 140 tion to anatomical structure the above-mentioned days. When analysing individual fruit development processes were connected mainly with the growth stages it was found that C. mas was slightly more ad- of the mesocarp. According to the results given by vanced in development in relation to C. officinalis. An- KANIEWSKI & HAUSBRANDT (1968), the growth of the thesis was observed in both investigated species with fleshy pericarp in C.