Citation: Kordyum, E.; Polishchuk,
O.; Akimov, Y.; Brykov, V.
Photosynthetic Apparatus of
Hydrocharis morsus-ranae in Different
Solar Lighting. Plants 2022, 11, 2658.
https://doi.org/10.3390/
plants11192658
Academic Editors: Bernhard
Huchzermeyer and Hans-Werner
Koyro
Received: 18 August 2022
Accepted: 3 October 2022
Published: 10 October 2022
Publisher’s Note: MDPI stays neutral
with regard to jurisdictional claims in
published maps and institutional affil-
iations.
Copyright: © 2022 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
plants
Article
Photosynthetic Apparatus of Hydrocharis morsus-ranae in
Different Solar Lighting
Elizabeth Kordyum
1
, Oleksandr Polishchuk
1
, Yuri Akimov
1
and Vasyl Brykov
1,2,
*
1
Department of Cell Biology and Anatomy, M.G. Kholodny Institute of Botany, National Academy of Sciences
of Ukraine, 2 Tereschenkivska Str., 01024 Kyiv, Ukraine
2
Department of Experimental Plant Biology, Faculty of Science, Charles University, Viniˇ cná 5,
128 00 Prague, Czech Republic
* Correspondence: vbrykov@gmail.com
Abstract: Hydrocharis morsus-ranae is a free-floating species growing in lakes and slow-flowing rivers
near the shore in Europe and Western Asia, and as an invasive plant in the USA and Canada. Light-
requiring plants of this species can also grow in the shade, up to about 30% of full sunlight. In this
paper we present the data about the photosynthetic apparatus of sunny and shady H. morsus-ranae
plants grown in the sun and in the shade in nature. Methods of light and transmission electron
microscopy, biochemistry, chlorophyll fluorescence induction as well as the principal component
analysis were used. It was found that leaves of plants growing in shade differed from those in the
sun with such traits as thickness of a blade, palisade and spongy parenchyma, ultrastructure of
chloroplasts, and quantum efficiency of photosynthetic electron transport, the content of chlorophylls
and carotenoids, anthocyanins and phenilpropanoids. By these traits, H. morsus-ranae shady plants
are similar with shade-bearing plants that indicates their adaptation to light intensity lowering. The
ordination plots (PCA) suggested a clear structural and functional shift of plants growing in different
lighting showing relationship to light changes in the natural environment. Thus, our results displayed
the high phenotypic plasticity of the H. morsus-ranae photosynthetic apparatus, which ensures its
acclimation to changing light environment and wide distribution of this species.
Keywords: acclimation; anthocyanin; chloroplast; chlorophyll induction; granum; pigment complex;
plasticity; principal component analysis; shade; sunlight
1. Introduction
Hydrocharis morsus-ranae L. are monocotyledonous, dioecious, stolon-rosette free-
floating aquatic plants (hydrophytes) of lakes, ponds and slow-flowing rivers, growing near
the shore [1]. Flowers are white, reaching almost 2 cm in diameter. Male (staminate) flowers
are usually gathered in inflorescences of three, female (pistillate) flowers are solitary. Heart
shaped leaves with the spongy lower side and long petioles are collected in the rosette and
float on the water surface. The color of leaves is usually a light or pure green or becoming
brownish black at senescence. It can vary depending on the level of sunlight or shade, and
plant age. In autumn, thin stolons with large buds (turions) at the ends appear. Turions
fall to the bottom of the water body, where they remain until spring. In spring, turions
develop new plants [2–5]. Micromorphology of vegetative and generative organs were
described by Cutter and Feldman [6], Seago et al. [7], Tsyrenova et al. [8], Efremov et al. [9].
H. morsus-ranae is native for Europe and Western Asia, and it widely naturalized outside its
native range in the USA and Canada as an invasive species because of its significant ability
to overgrow areas in a short time by vegetative and seed propagation [4,10]. In Ukraine,
H. morsus-ranae is usual in the Polissya (northern forest) and Forest-Steppe physiographic
regions, sporadically—in Steppe and on the Zakarpatska plain [11].
Plants 2022, 11, 2658. https://doi.org/10.3390/plants11192658 https://www.mdpi.com/journal/plants