Micromorphology and anatomy of the flower of Zephyranthes candida (Amaryllidaceae)

  • O. S. Fishchuk Lesya Ukrainka Eastern European National University
Keywords: flower anatomy; septal nectary; ovary; gynoecium; vascular bundle; vertical zonality; flower morphology


The use of morphological features of flowers in the taxonomy of plants is becoming increasingly important. The structure of the Zephyranthes candida (Lindl.) Herb. flowers on permanent cross-sectional and longitudinal sections was studied using a light microscope. The genus Zephyranthes belongs to the subtribe Hippeastrinae Walp. tribe Hippeastreae Sweet., family Amaryllidaceae s.l. Microscopic studies of the flower are considered as a tool to identify hitherto unknown structural adaptations of plants to specialized pollination methods and to elucidate the first stages of fruit morphogenesis, as many features of the fruit appear at the flower stage. The morphometric parameters, morphology, anatomy, and vascular anatomy of the ovary were described by using the flower’s transverse sections. Ten flowers of Z. candida were sectioned using standard methods of Paraplast embedding and serial sectioning at 20 μm thickness. Sections were stained with Safranin and Astra Blau and mounted in Eukitt. It was found that in the studied species the tepals have multi-bundle traces of 10–12 leading bundles. We consider the gynoecium of the studied species to be eusincarpous. The vascular system of the inferior ovary consists of three dorsal and three septal bundles, paired ventral bundles of carpels, which form ovule traces. For the first time, the presence of the following gynoecium zones was detected: a synascidiate structural zone with a height of about 360 μm and a fertile symplicate structural zone with a height of about 1560 μm and a hemisymplicate zone of 480 μm. Septal nectaries appear in the hemisymplicate zone and open with nectary split at the base of the style, the total height of the septal nectary is 760 μm. The ovary roof is 280 μm. Bifurcated dorsal and septal bundles of carpels have been identified, which can be considered as adaptations of the early stages of fruit morphogenesis to opening. Anatomical features of the ovary of Z. candida are numerous vascular bundles in the pericarpium, non-lignified endocarp at the flower stage, we consider as adaptations to the formation of juicy fruit. New data on the anatomical structure of the flower are a significant addition to the information on antecological and post-anthetic features of the studied species. Also, these data can be used in the construction of parsimony branches of the family Amaryllidaceae.


Afroz, S., Rahman, M., & Hassan, M. (2018). Taxonomy and reproductive biology of the genus Zephyranthes Herb. (Liliaceae) in Bangladesh. Bangladesh Journal of Plant Taxonomy, 25(1), 57–69.

Ao, C., Wang, L. Y., Sun, H., Lin, J. T., & Chen, Y. C. C. C. (2016). Megasporogenesis and megagametogenesis in Zephyranthes candida (Amaryllidaceae), with special notes on the behavior of the synergids, the central cell and the antipodal cells. Phyton, 56(1), 91–101.

Barykina, R. P., Veselova, T. D., Deviatov, A. G., Djalilova, H. H., Iljina, G. M., & Chubatova, N. V. (2004). Spravochnik po botanicheskoy mikrotehnike [Handbook of botanical microtechniques]. Izdatelstvo Moskovskogo Universiteta, Moscow (in Russian).

Büneker, H. M., & Bastian, R. E. (2018). Taxonomic novelties in Southern Brazilian Amaryllidaceae – II: Zephyranthes comunelloi a new species from Santa Catarina; and lectotypification of Zephyranthes mesochloa Herb. ex Lindl. Balduinia, 62, 1–8.

Centeno‐Betanzos, L. Y., Reyes‐Chilpa, R., Pigni, N. B., Jankowski, C. K., Torras-Claveria, L., & Bastida, J. (2021). Plants of the ‘Libellus de medicinalibus indorum herbis’ from Mexico 1552, Zephyranthes fosteri (Amaryllidaceae) alkaloids. Chemistry and Biodiversity, 18, e2000834.

Chase, M. W., Christenhusz, M. J. M., Fay, M. F., Byng, J. W., Judd, W. S., Soltis, D. E., Mabberley, D. J., Sennikov, A. N., Soltis, P. S., & Stevens, P. F. (2016). The angiosperm phylogeny group. An update of the angiosperm phylogeny group classification for the orders and families of flowering plants APG IV. Botanical Journal of the Linnean Society, 181, 1–20.

Chase, M. W., Reveal, J. L., & Fay, M. F. (2009). A subfamilial classification for the expanded asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae. Botanical Journal of the Linnean Society, 161(2), 132–136.

Chengqi, A. (2019). The endosperm development and the variations of structures of embryo sacs: Unravelling the low fecundity of Zephyranthes candida (Amaryllidaceae). Plant Biosystems, 153(5), 673–678.

Dash, C. K., Rahman, M. O., & Sultana, S. S. (2020). Karyological investigation on three Zephyranthes species and its taxonomic significance. Cytologia, 85(2), 163–168.

Daumann, E. (1970). Das Blütennektarium der Monocotyledonen unter besonderer Berücksichtigung seiner systematischen und phylogenetischen. Bedeutung Feddes Repertorium, 80(7–8), 463–590.

Fishchuk, O. (2021). Comparative flower morphology in Hippeastrum striatum (Lam.) H. E. Moore. (Amaryllidaceae). Ukrainian Journal of Ecology, 11(1), 273–278.

Francisco, L. (2017). A small flower with expansive energy: A proving of Zephyranthes rosea. Homoeopathic Links, 30(2), 112–113.

García, N., Meerow, A. W., Arroyo-Leuenberger, S., Oliveira, R. S., Dutilh, J. H., Soltis, P. S., & Judd, W. S. (2019). Generic classification of Amaryllidaceae tribe Hippeastreae. Taxon, 68(3), 425–612.

Katoch, D., & Singh, B. (2015). Phytochemistry and pharmacology of genus Zephyranthes. Medicinal and Aromatic Plants, 4(4), 212.

Kohelová, E., Maříková, J., Korábečný, J., Hulcová, D., Kučera, T., Jun, D., Chlebek, J., Jenčo, J., Šafratová, M., Hrabinová, M., Ritomská, A., Malaník, M., Peřinová, R., Breiterová, K., Kuneš, J., Nováková, L., Opletal, L., & Cahlíková, L. (2021). Alkaloids of Zephyranthes citrina (Amaryllidaceae) and their implication to Alzheimer’s disease: Isolation, structural elucidation and biological activity. Bioorganic Chemistry, 107, 104567.

Leinfellner, W. (1950). Der Bauplan des Syncarpen Gynoeceums. Oesterreichische. Botanische Zeitschrift, 97(3–5), 403–436.

Meerow, A. W., & Snijman, D. A. (1998). Amaryllidaceae. In: Kubitzki, K., Huber, H., Rudall, P. J., Stevens, P. S., & Studzel, T. (Ed.). The families and genera of vascular plants. III. Flowering plants: Monocotyledons: Lilianae (except Orchidaceae). Springer, Berlin. Pр. 83–110.

Meerow, A. W., Francisco-Ortega, J., & Schnell, R. J. (2006). Phylogenetic relationships and biogeography within the Eurasian clade of Amaryllidaceae based on plastid ndhF and nrDNA ITS sequences: Lineage sorting in a reticulate area? Systematic Botany, 31(1), 42–60.

Mokni, R. E., & Khalifa, K. H. (2020). More new geophytes for Tunisian and North African alien flora. Flora Mediterranea, 30, 185–196.

Murphy, P. J., Tibble-Howlings, J., Kowalczyk, R. M., & Stevens, K. (2020). Synthesis of zephycandidine A from haemanthamine. Tetrahedron Letters, 61(16), 151785.

Nguyen, K. V., Ho, D. V., Le, N. T., Phan, K. V., Heinämäki, J., Raal, A., & Nguyen, H. T. (2020). Flavonoids and alkaloids from the rhizomes of Zephyranthes ajax Hort. and their cytotoxicity. Scientific Reports, 10, 22193.

Nuraliev, M. S., & Sokoloff, D. D. (2014). Vaskulyarnaya anatomiya tsvetka kak istochnik informatsii ob evolyutsii pokryitosemennyih rasteniy: Istoriya izucheniya i sovremennyie predstavleniya [Floral vascular anatomy as a source of information on evolution of Angiosperms: History of study and current views]. Botanicheskiy Zhurnal, 99(2), 129–158 (in Russian).

Odinczova, A. (2013). Dva osnovnyh typy septalnyh nektarnykiv odnodolnyh [Two main types of septal nectaries in monocotyledons]. Visnyk Lvivskogo Universytetu, Seriya Biologichna, 61, 41–50 (in Ukrainian).

Odintsova, A., & Fishchuk, O. (2017). The flower morphology in three Convallariaceae species with various attractive traits. Acta Agrobotanica, 70(1), 1705–1719.

Shamrov, I. I. (2010). The peculiarities of syncarpous gynoecium formation in some monocotyledonous plants. Botanical Journal, 95(8), 1041–1070.

Smets, E. F., Ronse, De Craene, L. P., Caris, P., & Rudall, P. (2000). Floral nectaries in monocotyledons: Distribution and evolution. Monocots II: Systematics and Evolution. CSIRO, Melbourne. Pp. 230–240.

Spurrier, M. A., Smith, G. L., Flagg, R. O., & Serna, A. E. (2015). A new species of Zephyranthes (Amaryllidaceae) from Mexico. Novon, 24(3), 289–295.

Syeed, R., Mujib, A., Malik, M. Q., Mamgain, J., Ejaz, B., Gulzar, B., & Zafar, N. (2021). Mass propagation through direct and indirect organogenesis in three species of genus Zephyranthes and ploidy assessment of regenerants through flow cytometry. Molecular Biology Reports, 48, 513–526.

Takhtajan, A. (2009). Flowering plants. Springer, Dordrecht.

Torres-Morán, M. I., Velasco, P., Almaraz-Abarca, N., Anaya- Covarrubias, J. Y., & Velasco-Ramírez, A. (2019). Genetic structure of Zephyranthes fosteri, specie with ornamental and medicinal potential in Mexico. BIOtecnia, 21(2), 5–10.

Wu, B. W., & Liu, Y. T. (2017). First report of Hippeastrum chlorotic ringspot virus infecting Zephyranthes candida in China. Plant Disease, 101(11), 1960–1960.

Zhan, G., Liu, J., Zhou, J., Sun, B., Aisa, H. A., & Yao, G. (2017). Amaryllidaceae alkaloids with new framework types from Zephyranthes candida as potent acetylcholinesterase inhibitors. European Journal of Medicinal Chemistry, 127, 771–780.

Zhan, G., Qu, X., Liu, J., Tong, Q., Zhou, J., Sun, B., & Yao, G. (2016). Zephycandidine A, the first naturally occurring imidazo[1,2-f] phenanthridine alkaloid from Zephyranthes candida, exhibits significant anti-tumor and anti-acetylcholinesterase activities. Scientific Reports, 6, 33990.

Zhan, G., Zhou, J., Liu, R., Liu, T., Guo, G., Wang, J., Xiang, M., Xue, Y., Luo, Z., Zhang, Y., & Yao, G. (2016). Galanthamine, plicamine, and secoplicamine alkaloids from Zephyranthes candida and their anti-acetylcholinesterase and anti-inflammatory activities. Journal of Natural Products, 79(4), 760–766.

Zhan, G., Zhou, J., Liu, T., Zheng, G., Aisa, H. A., & Yao, G. (2016). Flavans with potential anti-inflammatory activities from Zephyranthes candida. Bioorganic and Medicinal Chemistry Letters, 26(24), 5967–5970.

Zhang, Y., Qiao, L., & Wang, J. (2018). Effects of leaf area index and degree of canopy cover of green turf and ground cover plants on rainwater interception. Nature Environment and Pollution Technology, 17(2), 563–568.

How to Cite
Fishchuk, O. S. (2021). Micromorphology and anatomy of the flower of Zephyranthes candida (Amaryllidaceae) . Regulatory Mechanisms in Biosystems, 12(2), 192–198. https://doi.org/10.15421/022127