Modification of the epicuticular waxes of plant leaves due to increased sunlight intensity

Keywords: elm; cuticular wax; hydrocarbon composition; steppe climate; metabolic adaptation.


Climatic changes observed around the world in recent years are associated with an increase in the solar radiation intensity and temperature and reduction in the humidity. Fluctuations of environmental factors significantly change the conditions for the existence of plants, which dictates the need for adaptive reactions of plant organisms at the different levels of their organization. Such dangerous processes as excessive heating of the surface of plant leaves and water loss can be prevented by the formation of a cuticle, which is a complex composition consisting of cutin and the soluble intracuticular and epicuticular waxes. We suggested that the structure, component composition and properties of the cuticle of trees undergo adaptive changes due to microclimatic conditions in different parts of the tree crown. The study was aimed at the identification and evaluation of light-induced differences in the accumulation and composition of leaf epicuticular waxes of Ulmus trees (native U. minor Mill. and alien U. pumila L.), and was conducted in 2018–2019 in Dnipro city located in the steppe zone of Ukraine. Analysis of the waxes’ chloroform extracts was carried out using GC Shimadzu 2010 PLUS equipped with a flame ionization detector and capillary column SP-2560. The highest amount of epicuticular waxes (12.23 ± 0.39 µg/cm2) was on the sunlit leaves of U. pumila, and wax deposits on the sunned leaves exceeded twice those on the shaded leaves in both U. minor and U. pumila. Long-chain hydrocarbons detected in the epicuticular waxes of both elm species were represented by free fatty acids, aldehydes, alcohols, and n-alkanes in various ratios. In the epicuticular waxes of U. minor, fatty acids dominated both on shaded and sunned leaves, while alkanes together with alcohols were the main components in U. pumila waxes, especially on the sunlit leaves. According to our results, local high illumination of leaves in the crown of both elm species caused increase in share of long-chain alkanes (1.2–1.9 times), but simultaneous reduction of the content of free fatty acids (1.5–16.8 times) in the epicuticular waxes’ composition. General patterns of the leaf epicuticular waxes’ modification due to increased solar radiation and air temperature can indicate the adaptive metabolic responses of woody plants to changing climatic conditions.


Bohinc, T., Markovič, D., & Trdan, S. (2014). Leaf epicuticular wax as a factor of antixenotic resistance of cabbage to cabbage flea beetles and cabbage stink bugs attack. Acta Agriculturae Scandinavica, Section B – Soil & Plant Science, 64(6), 493–500.

Bringe, K., Schumacher, C. F., Schmitz-Eiberger, M., Steiner, U., & Oerke, E. C. (2006). Ontogenetic variation in chemical and physical characteristics of adaxial apple leaf surfaces. Phytochemistry, 67(2), 161–170.

Brygadyrenko, V. V. (2016). Effect of canopy density on litter invertebrate community structure in pine forests. Ekológia (Bratislava), 35(1), 90–102.

Brygadyrenko, V. V., & Nazimov, S. S. (2014). Nutrition of Opatrum sabulosum (Coleoptera, Tenebrionidae) when fed on leaves of trees, shrubs and liana plants in the conditions of a laboratory experiment. Baltic Journal of Coleopterology, 14(1), 59–72.

Buschhaus, C., & Jetter, R. (2011). Composition differences between epicuticular and intracuticular wax substructures: How do plants seal their epidermal surfaces? Journal of Experimental Botany, 62(3), 841–853.

Buschhaus, C., Herz, H., & Jetter, R. (2007). Chemical composition of the epicuticular and intracuticular wax layers on adaxial sides of Rosa canina leaves. Annals of Botany, 100(6), 1557–1564.

Bussotti, F., Pollastrini, M., Holland, V., & Brüggemann, W. (2015). Functional traits and adaptive capacity of European forests to climate change. Environmental and Experimental Botany, 111(3), 91–113.

Didur, O., Kulbachko, Y., & Maltsev, Y. (2018). Impact of tropho-metabolic activity of earthworms (Lumbricidae) on distribution of soil algae within Acer platanoides L. plantation in recultivated territories of Western Donbass (Ukraine). Ukrainian Journal of Ecology, 8(2), 18–23.

Didur, O., Kulbachko, Y., Ovchynnykova, Y., Pokhylenko, A., & Lykholat, T. (2019). Zoogenic mechanisms of ecological rehabilitation of urban soils of the park zone of megapolis: Earthworms and soil buffer capacity. Journal of Environmental Research, Engineering and Management, 75(1), 24–33.

Domínguez, E., Cuartero, J., & Heredia, A. (2011). An overview on plant cuticle biomechanics. Plant Science, 181(2), 77–84.

Engelsdorf, T., Will, C., Hofmann, J., Schmitt, C., Merritt, B. B., Rieger, L., Frenger, M. S., Marschall, A., Franke, R. B., Pattathil, S., & Voll, L. M. (2017). Cell wall composition and penetration resistance against the fungal pathogen Colletotrichum higginsianum are affected by impaired starch turnover in Arabidopsis mutants. Journal of Experimental Botany, 68(3), 701–713.

González, A., & Ayerbe, L. (2010). Effect of terminal water stress on leaf epicuticular wax load, residual transpiration and grain yield in barley. Euphytica, 172(3), 341–349.

Grant, R. H., Heisler, G. M., Gao, W., & Jenks, M. (2003). Ultraviolet leaf reflectance of common urban trees and the prediction of reflectance from leaf surface characteristics. Agricultural and Forest Meteorology, 120, 127–139.

Guo, Y., Guo, N., He, Y., & Gao, J. (2015). Cuticular waxes in alpine meadow plants: Climate effect inferred from latitude gradient in Qinghai-Tibetan Plateau. Ecology and Evolution, 5(18), 3954–3968.

Guzmán-Delgado, P., Graça, J., Cabral, V., Gil, L., & Fernández, V. (2016). The presence of cutan limits the interpretation of cuticular chemistry and structure: Ficus elastica leaf as an example. Physiology Plantarum, 157 (2), 205–220.

Hansjakob, A., Bischof, S., Bringmann, G., Riederer, M., & Hildebrandt, U. (2010). Very-long-chain aldehydes promote in vitro prepenetration processes of Blumeria graminis in a dose- and chain length-dependent manner. New Phytologist, 188(4), 1039–1054.

Jetter, R., & Riederer, M. (2016). Localization of the transpiration barrier in the epi- and intracuticular waxes of eight plant species: Water transport resistances are associated with fatty acyl rather than alicyclic components. Plant Physiology, 170, 921–934.

Khromykh, N. O., Lykholat, Y. V., Kovalenko, I. M., Kabar, A. M., Didur, O. O., & Nedzvetska, M. I. (2018a). Variability of the antioxidant properties of Berberis fruits depending on the plant species and conditions of habitat. Regulatory Mechanisms in Biosystems, 9(1), 56–61.

Kim, K. S., Park, S. H., & Jenks, M. A. (2007). Changes in leaf cuticular waxes of sesame (Sesamum indicum L.) plants exposed to water deficit. Journal of Plant Physiology, 64(9), 1134–1143.

Kim, K. W., Ahn, J. J., & Lee, J. H. (2009). Micromorphology of epicuticular wax structures of the garden strawberry leaves by electron microscopy: Syntopism and polymorphism. Micron, 40(3), 327–334.

Klymenko, A., Kovalenko, І., Lykholat, Y., Khromykh, N., Didur, O., & Alekseeva, A. (2017). The integral assessment of the rare plant populations. Ukrainian Journal of Ecology, 7(2), 201–209.

Koch, K., & Ensikat, H. J. (2008). The hydrophobic coatings of plant surfaces: Epicuticular wax crystals and their morphologies, crystallinity and molecular self-assembly. Micron, 39(7), 759–772.

Kosma, D. K., Bourdenx, B., Bernard, A., Parsons, E. P., Lü, S., Joubès, J., & Jenks, M. A. (2009). The impact of water deficiency on leaf cuticle lipids of Arabidopsis. Plant Physiology, 151, 1918–1929.

Kunst, L., & Samuels, L. (2009). Plant cuticles shine: Advances in wax biosynthesis and export. Current Opinion in Plant Biology, 12, 721–727.

Lykholat, T. Y., Lykholat, O. A., Marenkov, O. M., Kulbachko, Y. L., Kovalenko, I. M., & Didur, O. O. (2019). Xeneostrogenes influence on cholinergic regulation in female rats of different age. Ukrainian Journal of Ecology, 9(1), 240–243.

Lykholat, T., Lykholat, O., & Antonyuk, S. (2016). Immunohistochemical and biochemical analysis of mammary gland tumours of different age patients. T︠s︡itologii︠a︡ i Genetika, 50(1), 40–51.

Lykholat, Y. V., Khromykh, N. O., Pirko, Y. V., Alexeyeva, A. A., Pastukhova, N. L., & Blume, Y. B. (2018a). Epicuticular wax composition of leaves of Tilia L. trees as a marker of adaptation to the climatic conditions of the Steppe Dnieper. Cytology and Genetics, 52(5), 323–330.

Lykholat, Y., Khromykh, N., Alexeyeva, A., Serga, O., Yakubenko, B., & Grigoryuk, I. (2017). Status of stomata and cuticular wax composition of the leaves of linden (Tilia tomentosa Moench.) under conditions of illumination and shading. Introduction of Plants, 2(74), 89–97 (in Ukrainian).

Lykholat, Y., Khromykh, N., Didur, O., Alexeyeva, A., Lykholat, T., & Davydov, V. (2018b). Modeling the invasiveness of Ulmus pumila in urban ecosystems in conditions of climate change. Regulatory Mechanisms in Biosystems, 9(2), 161–166.

Muller, C., & Riederer, M. (2005). Plant surface properties in chemical ecology. Journal of Chemical Ecology, 31(11), 2621–2651.

Nazarenko, M. M., & Lykholat Y. V. (2018). Influence of relief conditions on plant growth and development. Bulletin of the University of Dnepropetrovsk, Geology, Geography, 26(1), 143–149.

Nazarenko, M., Lykholat, Y., Grigoryuk, I., & Khromykh, N. (2018). Optimal doses and concentrations of mutagens for winter wheat breeding purposes. Part I. Grain productivity. Journal of Central European Agriculture, 19(1), 194–205.

Neinhuis, C., Koch, K., & Barthlott, W. (2001). Movement and regeneration of epicuticular waxes through plant cuticles. Planta, 213(3), 427–434.

Nobusawa, T., Okushima, Y., Nagata, N., Kojima, M., Sakakibara, H., & Umeda, M. (2013). Synthesis of very-long-chain fatty acids in the epidermis controls plant organ growth by restricting cell proliferation. PLoS Biology, 11(4), e1001531.

Pertseva, T., Lykholat, O., & Gurzhiy, O. (2012). Influence of tiotropium bromide (TB) and carbocysteine (C) on mucociliary clearance (MCC) in patients with COPD. European Respiratory Journal, 40(56), 3466.

Pokhylenko, A., Lykholat, O., Didur, O., Kulbachko, Y., & Lykholat, T. (2019). Morphological variability of Rossiulus kessleri (Diplopoda, Julida) from different biotopes within Steppe Zone of Ukraine. Ukrainian Journal of Ecology, 9(1), 176–182.

Racovita, R. C., & Jetter, R. (2016). Composition of the epicuticular waxes coating the adaxial side of Phyllostachys aurea leaves: Identification of very-long-chain primary amides. Phytochemistry, 130, 252–261.

Ramirez-Valiente, J. A., Koehler, K., & Cavender-Bares, J. (2015). Climatic origins predict variations in photo protective leaf pigments in response to drought and law temperature in live oaks (Quercus series virentes). Tree Physiology, 35(1), 521–534.

Reina-Pinto, J. J., & Yephremov, A. (2009). Surface lipids and plant defenses. Plant Physiology and Biochemistry, 47(6), 540–549.

Ringelmann, A., Riedel, M., Riederer, M., & Hildebrandt, U. (2009). Two sides of a leaf blade: Blumeria graminis needs chemical cues in cuticular waxes of Lolium perenne for germination and differentiation. Planta, 230(1), 95–105.

Shcherbyna, R. O., Danilchenko, D. M., Parchenko, V. V., Panasenko, O. I., Knysh, E. H., Khromykh, N. O., & Lykholat, Y. V. (2017). Studying of 2-((5-R-4-R1-4H-1,2,4-triazole-3-yl)thio)acetic acid salts influence on growth and progress of blackberries (Kiowa variety) propagules. Research Journal of Pharmaceutical, Biological and Chemical Science, 8, 975–979.

Shepherd, T., & Griffiths, W. D. (2006). The effects of stress on plant cuticular waxes. New Phytologist, 171(3), 469–499.

Tanaka, H., & Machida, Y. (2006). The cuticle and cellular interactions. In: Riederer, M., & Müller, C. (Eds.). Biology of the plant cuticle. Blackwell Publishing, Oxford. Pp. 312–333.

Van Maarseveen, C., & Jetter, R. (2009). Composition of the epicuticular and intracuticular wax layers on Kalanchoe daigremontiana (Hamet et Perr. de la Bathie) leaves. Phytochemistry, 70(7), 899–906.

Wen, M., Buschhaus, C., & Jetter, R. (2006). Nanotubules on plant surfaces: Chemical composition of epicuticular wax crystals on needles of Taxus baccata L. Phytochemistry, 67(16), 1808–1817.

Yeats, T. H., & Rose, J. K. (2013). The formation and function of plant cuticles. Plant Physiology, 163(1), 5–20.

Zeisler, V., & Schreiber, L. (2016). Epicuticular wax on cherry laurel (Prunus laurocerasus) leaves does not constitute the cuticular transpiration barrier. Planta, 243(1), 65–81.