Ginkgo biloba L. is a valuable relict species that combines high ornamental appeal, pharmacological importance, and resistance to a range of adverse environmental factors. In modern landscaping, more than 300 cultivars of G. biloba are used, differing in their morphological and ornamental characteristics. However, the use of the cultivar diversity of G. biloba in urban plantings requires consideration not only of their decorative value but also of their resistance to unfavorable environmental factors, primarily drought. The relevance of this study arises from the need to identify G. biloba cultivars most suitable for cultivation under the arid conditions of urbanized areas, as well as to investigate xeromorphic traits that reflect the adaptive potential of plants and may serve as criteria for their selection in urban greening. The aim of the study was to examine the morphometric characteristics of leaf blades and parameters of the stomatal apparatus in G. biloba plants and its various cultivars, and to identify genotypes with the highest expression of xeromorphic traits and adaptive capacity to water deficit. Morphometric measurements of the stomatal apparatus were carried out using the epidermal imprints method. The main morphometric parameters of leaves, as well as quantitative and morphometric characteristics of the stomatal apparatus components — including stomatal density, stomatal area, stomatal pore area, stomatal index, and xeromorphy index — were determined. Significant cultivar-specific differences were identified in a complex of leaf morphological and anatomical traits, which may reflect varying levels of xeromorphy and potential drought tolerance. It was established that the leaf surface area of G. biloba cultivars varies widely, ranging from 2.2 to 35.2 cm2, with the lowest values recorded in the cultivars ‘Chotek’ (needle-like leaves), ‘Saratoga’, and ‘Mariken’, being 4.1—16.0 times lower than in the control. This indicates a high degree of xeromorphic expression in these cultivars based on leaf morphometric characteristics. Based on the complex of xeromorphic characteristics of the stomatal apparatus (minimum stomatal area and high values of the stomatal index, and xeromorphy index), the cultivars ‘Mariken’, ‘Chotek’, ‘Troll’, and ‘Fastigiata’ were distinguished, indicating their considerable adaptive potential under conditions requiring optimized water use and suggesting their tolerance to moisture deficit. Given that these cultivars exhibit the broadest range of strongly expressed xeromorphic traits — including small leaf area and reduced dimensions of stomatal apparatus elements, along with high stomatal density, stomatal index, and xeromorphy index — their use should be prioritized when planning resilient urban green spaces.
Keywords: Ginkgo biloba L., stomatal apparatus, xeromorphic traits, stomatal denisty, stomatal index, xeromorphy index
Full text and supplemented materials
Free full text: PDFReferences
1. Crane, P.R. (2019). An evolutionary and cultural biography of ginkgo. Plants, People, Planet, 1(1), pp. 32-37. https://doi.org/10.1002/ppp3.7
2. Isah, T. (2015). Rethinking Ginkgo biloba L.: Medicinal uses and conservation. Pharmacog. Review, 9(18), pp. 140-148. https://doi.org/10.4103/0973-7847.162137
3. Laurain-Mattar, D. (2000). Cultivation of Ginkgo biloba on a large scale. Ginkgo biloba: medicinal and aromatic plants - industrial profiles: R. Hardman (Series ed.), T.A. van Beek (Volume ed.). Chapter 4. Publisher: Harwood Academic Publishers, pp. 63-79.
4. Del Tredici, P. (2000). The evolution, ecology, and cultivation of Ginkgo biloba. Ginkgo biloba. Australia, Canada, France, Germany, India, Japan, Luxembourg, Malaysia, The Netherlands, Singapore, Switzerland: Harwood academic publishers, pp. 7-23.
5. Ostudimov, A.O. & Guz, M.M. (2010). Peculiarities of ginkgo biloba seed reproduction. Scientific Bulletin of UNFU, 20(11), pp. 8-16 [in Ukrainian].
6. Ivaniuk, T.M., Kotiuk, L.A., Krasevych, N.O., Trofymenko, P.I. & Mykhaylovs'kyy, L.V. (2013). Botanichnyy sad Zhytomyrs'koho natsional'noho ahroekolohichnoho universytetu: inform.-dovid. Putivnyk: Zhytomyr: Zhytomyrs'kyy natsional'nyy ahroekolohichnyy universytet, pp. 149-150.
7. Lin, H.Y., Li, W.H., Lin, C.F., Wu, H.R. & Zhao, Y.P. (2022). International biological flora: Ginkgo biloba. J. Ecol., 110(4), pp. 951-982. https://doi.org/10.1111/1365-2745.13856
8. Kayser, O. & Quax, W.J. (Eds.) (2006). Medicinal plant biotechnology: from basic research to industrial applications. Chap. 21. Wiley-VCH, pp. 493-514. https://doi.org/10.1002/9783527619771
9. Leontiak, H.P. (2016, May). Hinkho dvolopateve - velyke maybutnie v Ukraini ta Moldovi. Perspektyvy rozvytku lisovoho ta sadovo-parkovoho hospodarstva. Materialy Vseukr. nauk.-prakt. konferentsii. Uman': VPTs «Vizavi» pp. 25-35 [in Ukrainian].
10. Davies, T. (2022). The (UK) national plant collection of Ginkgo biloba & Cultivars. Res. Int. Retrieved from https://www.npcginkgo.org/npcginkgo
11. ћmarda, P., Horov«, L., Kn«pek, O., Dieck, H., Dieck, M., Raыn«, K., HrubHk, P., OrlЩci, L., Papp, L., Vesel«, K., Veselъ, P. & Bureл, P. (2018). Multiple haploids, triploids, and tetraploids found in modernday "living fossil" Ginkgo biloba. Horticult. Res., 5(1), pp. 1-12. https://doi.org/10.1038/s41438-018-0055-9
12. Filipczak, J. (Ed.). (2013). Katalog rasteniy (derev'ia, kustarniki, mnogoletniki rekomendovannye Soiuzom Pol'skikh Pitomnikov). Warszawa, Polska: Agencja Promocji Zieleni Sp. z o.o. pp. 26-29.
13. Shumyk, M.I., Kliuienko, O.V., Korkulenko, O.M., Popil, N.I. & Ostapyuk, V.M. (2018). Ontomorphogenesis of summergreen (deciduous) species of the genus Rhododendron L. ex situ. Plant Int., 3, pp. 39-51. Res. Int. Retrieved from http://jnas.nbuv.gov.ua/article/UJRN-0001035561 [in Ukrainian].
14. Leshcheniuk, O.M. & Mazura M.Yu. (2021). Changes in the anatomical parameters of the leaves of Forsythia europaea Degen & Bald. caused by motor vehicle emissions. Sci. Bulletin of UNFU., 31(5), pp. 29-35. [in Ukrainian]. https://doi.org/10.36930/40310504
15. Vlasenko, N.O. (2025). Strategies for tree adaptation to urban environments. Sci. Bulletin of UNFU., 35 (1), pp. 9-15. [in Ukrainian]. https://doi.org/10.36930/40350101
16. Nuzhyna, N.V. & Palagecha, R.M. (2020). Anatomic features of leaf of relict plant species in connection with drought resistance. Pryrodnychyy al'manakh (biolohichni nauky), 28, pp. 75-84. [in Ukrainian]. https://doi.org/10.32999/ksu2524-0838/2020-28-7
17. Krugliak, Yu. (2018). Morphometric characteristics of stomata of plants Philadelphus L. genus in connection with their drought resistance. Biol. Syst., 10(2), pp. 193-197. https://doi.org/10.31861/biosystems2018.02.193
18. Shchyrova, Yu.V., Serbin, A.H. & Kartmazova, L.S. (2002). Morfoloho-anatomichne doslidzhennia lystkiv hinkho dvolopatevoho. Visnyk farmatsii, 4(32), pp. 19-22. Res. Int. Retrieved https://dspace.nuph.edu.ua/bitstream/123456789/1336/1/19-22(1).pdf [in Ukrainian].
19. Rymar, Yu.Yu., Pronina, O.V., Kiriziy, D.A., Duplii, V.P., Morgun, B.V. & Stasik, O.O. (2025). Characteristics of flag leaf stomata in relation to gas exchange rate and drought tolerance in related spring wheat species. FTzTol. rosl. genet., 57(1), pp. 64-82. [in Ukrainian]. https://doi.org/10.15407/frg2025.01.064
20. Xu, Z. & Zhou, G. (2008). Responses of leaf stomatal density to water status and its relationship with photosynthesis in a grass. J. Exp. Bot., 59(12), pp. 3317-3325. https://doi.org/10.1093/jxb/ern185
21. VolenHkov«, M. & Tich«, I. (2001). Insertion profiles in stomatal density and sizes in Nicotiana tabacum L. plantlets. Biol. Plant., 44(2), pp. 161-165. https://doi.org/10.1023/A:1017982619635
22. Chen, L.Q., Li, C.S., Chaloner, W.G., Beerling, D.J., Sun, Q.G., Collinson, M.E. & Mitchell, P.L. (2001). Assessing the potential for the stomatal characters of extant and fossil Ginkgo leaves to signal atmospheric CO2 change. American J. Bot., 88(7), pp. 1309-1315. https://doi.org/10.2307/3558342
23. Liu, C., Sack, L., Li, Y., Zhang, J., Yu, K., Zhang, Q., He, N. & Yu, G. (2023). Relationships of stomatal morphology to the environment across plant communities. Nature Comm., 14(1), Art. 6629. https://doi.org/10.1038/s41467-023-42136-2
24. Rymar, Yu.Yu., Pronina, O.V., Duplii, V.P. & Morgun, B.V. (2023). Peculiarities of stomata morphology in bread wheat. Faktory eksperymental'noi evoliutsii orhanizmiv, 32, pp. 120-124. [in Ukrainian]. https://doi.org/10.7124/FEEO.v32.1547
25. Demchenko, N., Futorna, O., Badanina, V., Smirnov, O., Olshanskyi, I. & Taran, N.T. (2019). Stomata complexes of leaves of leaf-declining representatives of Magnoliaceae as a marcers of a thermoregulatory and microclimate-forming ability of plants. Ekol. Nauk., 1(24), pp. 149-159. [in Ukrainian]. https://doi.org/10.32846/2306-9716-2019-1-24-1-27
26. Kryvoruchko, A.P. & Bessonova, V.P. (2018). Anatomical leaves characteristics of Quercus rubra L. and Quercus robur L. and stand density. Ukrainian J. Ecol., 8(1), pp. 64-71. https://doi.org/10.15421/2018_188
27. Atramentova, L.O. & Utievs'ka, O.M. (2007). Biometriia. Ch. II. Porivniannia hrup i analiz zv'iazku. Kharkiv: Ranok [in Ukrainian].
28. Atramentova, L.O. & Utievs'ka, O.M. (2014). Statystyka dlia biolohiv. Kharkiv: NTMT [in Ukrainian].
29. Roth-Nebelsick, A. & Krause, M. (2023). The plant leaf: A biomimetic resource for multifunctional and economic design. Biomim., 8(2), pp. 1-32. https://doi.org/10.3390/biomimetics8020145
30. Kramp, R.E., Liancourt, P., Herberich, M.M., Saul, L., Weides, S., TielbШrger, K. & M«jekov«, M. (2022). Functional traits and their plasticity shift from tolerant to avoidant under extreme drought. Ecol., 103(12), pp. 1-8. https://doi.org/10.1002/ecy.3826
31. Nedukha, O.M. (2015). Klitynna obolonka roslyn i faktory seredovyshcha. Kyiv: Al'terpres. Res. Int. Retrieved https://www.botany.kiev.ua/doc/nedyx_2015.pdf [in Ukrainian].
32. Hetherington, A.M. & Woodward, F.I. (2003). The role of stomata in sensing and driving environmental change. Nature, 424, pp. 901-908. https://doi.org/10.1038/nature01843
33. Lawson, T. (2009). Guard cell photosynthesis and stomatal function. New Phytol., 181(1), pp. 13-34. https://doi.org/10.1111/j.1469-8137.2008.02685.x
34. Strobel, D.W. & Sundberg, M.D. (1983). Stomatal density in leaves of various xerophytes: a preliminary study. J. Minnesota Academ. Sci., 49 (2), pp. 7-9. Res. Int. Retrieved https://digitalcommons.morris.umn.edu/cgi/viewcontent.cgi?article=1841 &context=jmas
35. Xi, J.J., Chen, H.Y., Bai, W.P., Yang, R.C., Yang, P.Z., Chen, R.J., Hu, T.M. & Wang, S.M. (2018). Sodium-related adaptations to drought: New insights from the xerophyte Zygophyllum xanthoxylum. Front. Plant Sci., 9, Art. 1678, pp. 1-15. https://doi.org/10.3389/fpls.2018.01678
36. Zhygalova, S.L. & Futorna, O.A. (2013). The micromorphological features of Gladiolus imbricatus L. (Iridaceae Juss.). Modern Phytomorphol., 3, pp. 273-280 [in Ukrainian].
37. Guan, Z.J., Zhang, S.B., Guan, K.Y., Li, S.Y. & Hu, H. (2011). Leaf anatomical structures of Paphiopedilum and Cypripedium and their adaptive significance. J. Plant Res., 124, pp. 289-298. https://doi.org/10.1007/s10265-010-0372-z