Phenotypic variability of plant leaves of Acer genus, introduced into steppe zone of Ukraine
Abstract
This paper deals with studying of the patterns of ecological adaptation of wood species of Acer L. genus during their introduction into steppe zone of Ukraine. Hydrothermal conditions of the growing season in steppe zone of Ukraine are particularly unfavorable for tree and shrubbery plantings, comprising both native and introduced species. In the course of plants’ introduction, adaptive changes occur; such changes represent the spectrum of phenotypic implementation of the definite genotype under the influence of new environmental conditions. Stress environment of the region of introduction leads to occurrence of a great variety of phenotypic forms, as the different variants of genotype implementation. Studying of phenotypic variability gives an opportunity to determine the capacity to adaptation of introduced species and ways of adaptive reactions in new conditions of living. Therefore, objective of the work consists in studying of the processes of differentiation of morphological characters in species of Acer genus introduced in the regions of steppe zone with varying intensity of hydrothermal factors. Studies were carried out in the central and south-eastern steppe regions, as well as in the south of steppe zone in the coastal and continental areas. Subjects of research were 9 species of maples, differing by their botanic and geographic origin and by the degree of drought resistance in the steppe zone of Ukraine. Patterns of variability of morphostructural characters of leaves were determined by the indicator of specific weight of leaves which was calculated as a ratio of weight of dry laminas to their area (mg/cm2). Following the results of study, it was found that adaptation of maples to xerothermic factors of the environment is connected with changing of the ratio of groups of character variation and their contribution into total sample. Direct relationship is established between the probability density of expression (phenotypic implementation) of the character in the definite range of variation and degree of adaptation of the plant. Typical feature of species with the most prominent xeromorphic characters (A. campestre, A. tataricum, A. monspessulanum) is gradual increase in both the mean value of one of the components of the leaves’ specific weight distribution curve and its contribution in the total sample with the increasing hydrothermal stress. In maple species with low stability, probability of the character display is actually the same for each segregate component of total sample in the different regions of steppe zone, which indicates that these species has no adaptive potential to be realized in the specific environment. Eco-climatic adaptation of mesophytic maple species is displayed in reduction of confidence intervals in the segregate components of the integral curve and reduction of total interval of variation of specific weight of leaves, i.е. it is aimed at decrease in phenotypic polymorphism of introduced species of Acer genus in the drought conditions of steppe zone of UkraineReferences
Aranda, I., Pardo, F., Gil, L., Pardos, J.A., 2004. Anatomical basis of the change in leaf mass per area and nitrogen investment with relative irradiance within the canopy of eight temperate tree species. Acta Oecol. 25(3), 187–195. >> doi.:10.1016/j.actao.2004.01.003
Bobyliov, Y.P., Brygadyrenko, V.V., Bulakhov, V.L., Gaichenko, V.A., Gasso, V.Y., Didukh, Y.P., Ivashov, A.V., Kucheriavyi, V.P., Maliovanyi, M.S., Mytsyk, L.P., Pakhomov, O.Y., Tsaryk, I.V., Shabanov, D.A., 2014. Ekologija [Ecology]. Folio, Kharkiv (in Ukrainian).
Borisjuk, V.A., Kljachenko, V.I., Tretjak, T.V., 1989. Anatomo-morfologicheskie i fiziologo-biohimicheskie testy pri izuche¬nii selekcionnyh materialov saharnoj svekly [Anatomical-morphological, physiological and biochemical tests in the study of sugar beet breeding materials]. Kolos, Moscow (in Russian).
Bosabalidis, A.M., Kofidis, G., 2002. Comparative effects of drought stress on leaf anatomy of two olive cultivars. Plant Sci. 163(2), 375–379. >> doi.:10.1016/S0168-9452(02)00135-8
Bulakhov, V.L., Emel'janov, I.G., Pakhomov, O.Y., 2003. Bioraznoobrazie kak funkcional'naja osnova jekosistem [Biodiversity as functional basis of ecosystems]. Vìsn. Dnìpropetr. Unìv. Ser. Bìol. Ekol. 11(1), 3–8.
Burda, R.I., 1991. Antropogennaja transformacija flory [Anthropogenic transformation of the flora]. Naukova Dumka, Kyiv (in Russian).
Chen, X.-H., Gao, Y.-B., Zhao, T.-T., Zhu, M.-J., Ci, H.-C., Xiao-Yan, S., 2010. Morphological variations of Caragana microphylla populations in the Xilingol steppe and their relationship with environmenral factors. Acta Ecol. Sin. 30(2), 50–55. >> doi.:10.1016/j.chnaes.2010.03.001
Choat, B., Ball, M.C., Luly, J.G., Donnelly, C.F., Holtum, J.A.M., 2006. Seasonal patterns of leaf gas exchange and water relations in dry rain forest trees of contrasting leaf phenology. Tree Physiol. 26, 657–664. >> doi.:10.1093/treephys/26.5.657
Fang, J., Wu, F., Yang, W., Zhang, J., Cai, H., 2012. Effects of drought on the growth and resource use efficiency of two endemic species in an arid ecotone. Acta Ecol. Sin. 32(4), 195–201. >>doi.:10.1016/j.chnaes.2012.05.001
Feng, R., Zhang, Y., Yu, W., Hu, W., Wu, J., Ji, R., Wang, H., Zhao, X., 2013. Analysis of the relationship between the spectral characteristics of maize canopy and leaf area index under drought stress. Acta Ecol. Sin. 33(6), 301–307. >> doi.:10.1016/j.chnaes.2013.09.001
Kapustjan, A., Zhuk, O., 2009. Morfogenetychna adaptyvna vidpovid' roslyn na stres [Morphogenetic adaptive response of plants to stress]. Visnyk Kyi'vs'kogo Nac. Univ. Serija «Introdukcija ta zberezhennja roslynnogo riznomanittja» 26, 79–81 (in Ukrainian).
Kohno, N.A., 1982. Klenі Ukraynі [Maples of Ukraine]. Naukova Dumka, Kyiv (in Russian).
Kordjum, J.L., 2001. Fenotypichna plastychnist' u roslyn: Stan problemy ta perspektyvy [Phenotypical plasticity in plants: Problems and prospects of state]. Materialy IX Z’i'zdu Ukr. Bot. Soc. Harkiv 180–181 (in Ukrainian).
Niinemets, Ü., Kull, K., 2003. Leaf structure vs. Nutrienr relationship vary with soil conditions in temperate shrubs and trees. Acta Oecol. 24(4), 209–219. >>doi.:10.1016/S1146-609X(03)00094-8
Pakhomov, O.Y., Brygadyrenko, V.V., 2005. Koncepcija systemy zahodiv z ohorony navkolyshn'ogo pryrodnogo seredovyshha Dnipropetrovs'koi' oblasti na 2005–2015 roky [Concept of system for actions on environment protection in Dnipropetrovsk region for 2005–2015]. Vìsn. Dnìpropetr. Unìv. Ser. Bìol. Ekol. 13(1), 213–225.
Pakhomov, O.Y., Gasso, V.Y., Goloborodko, K.K., Poljakov, M.V., Grycan, Y.I., Bulakhov, V.L., Brygadyrenko, V.V., Kljuchko, Z.F., Mezhzherin, S.V., Novicky, R.O., Pysanec, Y.M., Pljushh, I.G., Ponomarenko, O.L., Puchkov, O.V., Radchenko, V.G., 2011. Chervona knyga Dnipropetrovskoi oblasti. Tvarynnyj svit [The red book of Dnipropetrovsk region. Animals]. New Print, Dnipropetrovsk (in Ukrainian).
Raskatov, P.B., 1979. Еkologycheskaja anatomyja vegetatyvnіh organov derev'ev y kustarnykov [Ecological anatomy of vegetative organs of trees and shrubs]. Voronezh University Press, Voronezh (in Russian).
Rum’jankov, J., 2009. Anatomo-morfologichni oznaky kseromorfnosti lystkiv vydiv rodu Celtis L. v umovah Nacional'nogo dendroparku “Sofii'vka” NAN Ukrai'ny [Anatomical and morphological characteristics of leaves kseromorfity in species of Celtis L. in the National park "Sofiyivka", NAS Ukraine]. Introdukcija ta Zberezhennja Roslynnogo Riznomanittja 27, 143–144 (in Ukrainian).
Shavrov, L.A., 1961. Jekologo-geograficheskaja zakonomernost' strukturnoj izmenchivosti introducirovannyh rastenij [Ecological and geographical pattern of structural variability of introduced plants]. Botanicheskij Zh. 46(3), 328–336.
Slabbert, M.M., Krüger, G.H.J., 2014. Antioxidant enzyme activity, proline accumulation, leaf area and cell membrane stability in water stressed. South Afr. J. Bot. 95, 123–128.
Tooming, H.G., 1984. Еkologycheskye pryncypі maksymal'noj produktyvnosty posevov [Ecological principles of maximum productivity of crops]. Gidrometeoizdat, Leningrad (in Russian).
Waring, R.H., 1983. Estimating forest growth and efficiency in relation to canopy leaf area. Adv. Ecol. Res. 13, 327–354. >> doi.:10.1016/S0065-2504(08)60111-7
Zaitseva, I.A., 2004. Dynamika vodoobminnyh procesiv vydiv rodu Acer L. u zv’jazku z i'h posuhostijkistju [Waterprocesses dynamics for species of the Acer L. genus due to their drought]. Vìsn. Dnìpropetr. Unìv. Ser. Bìol. Ekol. 12(1), 54–62 (in Ukrainian).
Zaitseva, I.A., 2012. Izmenchivost' morfostrukturnyh priznakov list'ev drevesnyh introducentov roda Acer L. v stepnoj zone Ukrainy [Variability of morphostructural signs of leaves of Acer L. genus woody introducents in the steppe zone of Ukraine]. Visti Biosfernogo Zapovidnyka “Askanija-Nova” 14, 106–111 (in Russian).
Zajcev, G.N., 1983. Optimum i norma v introdukcii rastenij [Optimum and the norm in plant introduction]. Nauka, Moskow (in Russian).
Zhuang, L., Li, W.-H., Yuan, F., Gong, W.-C., Tian, Z.-P., 2010. Ecological adaptation characteristics of Populus euphratica and Tamarix ramosissima leaf microstructures in the lower reaces of Tarim River. Acta Ecol. Sin. 30(2), 62–66. >> doi.:10.1016/j.chnaes.2010.03.003