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Ботанический журнал, 2024, № 1

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Российская академия наук
БОТАНИЧЕСКИЙ 
ЖУРНАЛ
том 109        № 1        2024        Январь
Журнал основан в декабре 1916 г.
Выходит 12 раз в год
ISSN печатной версии 0006-8136
ISSN Online версии 2658-6339
Журнал издается под руководством
Отделения биологических наук РАН
Главный редактор
Л.В. Аверьянов
доктор биологических наук, профессор
Редакционная коллегия:
Афонина О.М. д.б.н., зам. глав. редактора; 
Сафронова И.Н. д.б.н., зам. глав. редактора;
Сытин А.К. д.б.н., зам. глав. редактора;
Шамров И.И. д.б.н., зам. глав. редактора
Баранова О.Г. д.б.н.; Волис С. (Китай) PhD, профессор; Герман А.Б. д.г.-м.н.;
Дарбаева Т.Е. (Казахстан) д.б.н.; Димеева Л.А. (Казахстан) д.б.н.;
Кузьмина М.Л. (Канада) PhD; Куликовский М.С. д.б.н.; Марков М.В. д.б.н.;
Михайлова Т.А. к.б.н.; Оскольский А.А. (ЮАР) д.б.н.; Палице З. (Чехия) PhD;
Паутов А.А. д.б.н., профессор; Пименов М.Г. д.б.н., профессор; 
Романов Р. Е. к.б.н.; Сенников А.Н. (Финляндия) к.б.н.; Соколов Д.Д. д.б.н., 
член-корр. РАН, профессор; Соколова И.В. к.б.н.; Тимонин А.К. д.б.н.; 
Тиходеева М.Ю. к.б.н.; Шнеер В.С. д.б.н.; Яковлев Г.П. д.б.н., профессор
Кессель Д.С. – отв. секретарь редколлегии,
Битюкова Н.В. – секретарь редколлегии
Зав. редакцией М.О. Набатова-Азовская
Ответственный редактор номера О.М. Афонина
E-mail: botzhurn@mail.ru, mari.nabatova-azovskaya@mail.ru
Москва
ФГБУ «Издательство «Наука»
© Российская академия наук, 2024 
© Редколлегия “Ботанический журнал”
(составитель), 2024


RUSSIAN ACADEMY OF SCIENCE 
BOTANICHESKII 

ZHURNAL
Volume 109
№ 1
MOSCOW  2024


Founders:
RUSSIAN ACADEMY OF SCIENCES BRANCH OF BIOLOGICAL 
SCIENCES RAS RUSSIAN BOTANICAL SOCIETY
BOTANICHESKII ZHURNAL
Periodicity 12 issues a year Founded in December 1916
Journal is published the algis of the Branch of Biological Sciences RAS
Editor-in-Chief
L. V. Averyanov, Doctor of Sciences (Biology)
EDITORIAL BOARD
O. M. Afonina (Deputy Editor-in-Chief, Doctor of Sciences (Biology), St. Petersburg, Russia),
I. N. Safronova (Deputy Editor-in-Chief, Doctor of Sciences (Biology), St. Petersburg, Russia),
I. I. Shamrov (Deputy Editor-in-Chief, Doctor of Sciences (Biology), St. Petersburg, Russia),
A. K. Sytin (Deputy Editor-in-Chief, Doctor of Sciences (Biology), St. Petersburg, Russia),
D. S. Kessel (Executive Secretary, St. Petersburg, Russia),
N. V. Bityukova (Secretary, St. Petersburg, Russia),
O. G. Baranova (Doctor of Sciences (Biology), St. Petersburg, Russia),
S. Volis (PhD, Kunming, China),
A. V. Herman (Doctor of Sciences (Geology and Mineralogy), Moscow, Russia),
T. E. Darbayeva (Doctor of Sciences (Biology), Uralsk, Kazakhstan),
L. A. Dimeyeva (Doctor of Sciences (Biology), Almaty, Kazakhstan),
M. L. Kuzmina (PhD, Guelph, Canada),
M.S. Kulikovskiy (Doctor of Sciences (Biology), Moscow, Russia),
M. V. Markov (Doctor of Sciences (Biology), Moscow, Russia),
T. A. Mikhaylova (Candidate of Sciences (Biology), St. Petersburg, Russia),
A. A. Oskolski (Doctor of Sciences (Biology), St. Petersburg, Russia; Johannesburg, RSA),
Z. Palice (PhD., Prùhonice, Czech Republic),
A. A. Pautov (Doctor of Sciences (Biology), St. Petersburg, Russia),
M. G. Pimenov (Doctor of Sciences (Biology), Moscow, Russia),
R. E. Romanov (Candidate of Sciences (Biology), St. Petersburg, Russia),
A. N. Sennikov (Candidate of Sciences (Biology), Helsinki, Finland),
D. D. Sokoloff (Doctor of Sciences (Biology), Moscow, Russia),
I. V. Sokolova (Candidate of Sciences (Biology), St. Petersburg, Russia),
M. J. Tikhodeeva (Candidate of Sciences (Biology), St. Petersburg, Russia)
A. C. Timonin (Doctor of Sciences (Biology), Moscow, Russia),
V. S. Shneyer (Doctor of Sciences (Biology), St. Petersburg, Russia),
G. P. Yakovlev (Doctor of Sciences (Biology), St. Petersburg, Russia)
Managing editor M. O. Nabatova-Azovskaya   Executive editor of the issue O. M. Afonina
E-mail: botzhurn@mail.ru, mari.nabatova-azovskaya@mail.ru
Moscow 2024
© Russian Academy of Sciences, 2024
© Compilation Editorial board
of “Botanicheskii Zhurnal”, 2024


СОДЕРЖАНИЕ
Том 109, Номер 1, 2024
ОБЗОРНЫЕ СТАТЬИ
Gamete structure and development in connection with fertilization in angiosperms 
I. I. Shamrov, G. M. Anisimova
3
ОРИГИНАЛЬНЫЕ СТАТЬИ
Low temperature-induced chloroplast relocation in mesophyll cells of Pinus sylvestris (Pinaceae):  
SBF SEM 3D reconstruction
N. K. Koteyeva, A. N. Ivanova, T. A. Borisenko, M. S. Tarasova, O. E. Mirgorodskaya, E. V. Voznesenskaya
29
СООБЩЕНИЯ
Распространение и особенности биологии Magadania olaensis (Apiaceae) 
О. А. Мочалова, М. Г. Хорева, Е. А. Андриянова
42
Полиморфизм геномной ДНК Myrica gale (Myricaceae) на территории Государственного  
природного заказника “Лебяжий” (южное побережье Финского залива)
О. А. Семичева, У. А. Галактионова, В. Н. Большаков, А. Э. Романович, М. Ю. Тиходеева, О. Н. Тиходеев
65
Сравнительный морфологический и генетический анализ популяций Corydalis bracteata Sensu Lato 
(Papaveraceae) из Южной Сибири
К. К. Рябова, И. Е. Ямских, Н. В. Степанов, М. Г. Куцев
81
ФЛОРИСТИЧЕСКИЕ НАХОДКИ
Artemisia verlotiorum (Asteraceae) – новый вид для флоры Азербайджана
Н. П. Мехтиева, К. К. Асадова, Н. Мурсал 
97
ХРОНИКА
III Всероссийская конференция с международным участием “Систематические и флористические 
исследования Северной Евразии” (к 95-летию со дня рождения профессора А.Г. Еленевского) 
(Москва, 19–21 октября 2023 г.)
В. П. Викторов, В. Н. Годин, Н. М. Ключникова, Н. Г. Куранова, С. К. Пятунина
101
 


CONTENTS
Vol. 109, Number 1, 2024
REVIEWS
Gamete structure and development in connection with fertilization in angiosperms 
I. I. Shamrov, G. M. Anisimova
3
ORIGINAL ARTICLES
Low temperature-induced chloroplast relocation in mesophyll cells of Pinus sylvestris (Pinaceae):  
SBF SEM 3D reconstruction
N. K. Koteyeva, A. N. Ivanova, T. A. Borisenko, M. S. Tarasova, O. E. Mirgorodskaya, E. V. Voznesenskaya
29
COMMUNICATIONS
Distribution and peculiarities of biology of Magadania olaënsis (Apiaceae) 
O. A. Mochalova, M. G. Khoreva, E. A. Andriyanova
42
Genomic DNA polymorphism in Myrica gale (Myricaceae) in the Lebyazhiy State Nature Reserve  
(the southern coast of the Gulf of Finland)
O. A. Semicheva, U. A. Galaktionova, V. N. Bolshakov, A. E. Romanovich, M. Yu. Tikhodeeva, O. N. Tikhodeyev
65
Comparative morphological and genetic analysis of Corydalis bracteata sensu lato (Papaveraceae) populations from 
Southern Siberia
K. K. Ryabova, I. Е. Yamskikh, N. V. Stepanov, M. G. Kutsev
81
FLORISTIC RECORDS
Artemisia verlotiorum (Asteraceae), a new species to the flora of Azerbaijan
N. P. Mehdiyeva, K. K. Asadova, N. Mursal 
97
CHRONICLES
III All-Russian Conference with international participation “Systematic and floristic research  
of Northern Eurasia” (on 95-year anniversary of Professor A.G. Elenevsky) (Moscow, October 19–21, 2023) 
V. P. Viktorov, V. N. Godin, N. M. Klyuchnikova, N. G. Kuranova, S. K. Pyatunina
101
 


БОТАНИЧЕСКИЙ ЖУРНАЛ, 2024, том 109, № 1, с. 5–30
 ОБЗОРНЫЕ СТАТЬИ 
GAMETE STRUCTURE AND DEVELOPMENT IN CONNECTION 
WITH FERTILIZATION IN ANGIOSPERMS
© 2024 г.      I. I. Shamrova,b,#, G. M. Anisimovab,##
aHerzen State Pedagogical University of Russia
Moika River Emb., 48, St. Petersburg, 191186, Russia
bKomarov Botanical Institute of RAS
Prof. Popov Str., 2, St. Petersburg, 197022, Russia
#e-mail: shamrov52@mail.ru
##e-mail: galina0353@mail.ru
Received 17.11.2023
Revised 03.12.2023
Accepted 05.12.2023
The paper analyzes data on the processes occurring before and during fertilization in flowering plants. 
At the gametophyte stage, the gametes are formed from haploid microspores and megaspores. They are 
sperms, egg and central cell. The fusion of male and female gametes occurs after the pollen tube enters 
any synergid. One sperm fuses with the egg cell, and another from the same pollen tube joins with the 
central cell.
The angiosperms are likely to have four types of fertilization. These types differ in the degree of 
completion of syngamy. Premitotic and postmitotic types are characterized by complete syngamy, and 
androgamic and gynandrogamic types are inherent in incomplete syngamy. In this case, the behavior 
of the sperm nuclei is of great significance. They, as a rule, combine with the nuclei of female gametes 
(premitotic and postmitotic types), but the sperm nuclei can remain independent (gynandrogamic type) 
or a female nucleus is replaced by a male one (androgamic type). The premitotic type of fertilization 
follows the way in which the gamete protoplasts and nuclei are united before the mitosis in the zygote 
nucleus. As for the postmitotic type, it is carried out on a completely different basis. It is possible that in 
angiosperms the chromosomes of sperm and egg cell do not unite during mitosis. They further divide 
independently, and a diploid set of chromosomes arises in a 2-celled embryo.
Keywords: male and female gametes, fertilization, syngamy, triple fusion, fertilization types, angiosperms
DOI: 10.31857/S0006813624010013, EDN: FFLKVQ
The gametes, represented in angiosperms 
of the zygote) (Gerassimova-Navashina, 1947, 
1957, 1960, 1969, 1990). Available information in 
the literature indicates that general principles of 
fertilization mechanisms in animals and flowering 
plants are more conserved than previously thought. 
Among them there are following aspects: structure 
by sperms and egg cells, unlike somatic cells, 
have a haploid set of chromosomes. The places 
of localization, the time of the appearance of 
gametes during ontogenesis, the features of differentiation and transformation, as well as some 
of gametes, cell–cell communication events between 
gametes as well as their physical interaction and fusion 
during fertilization (Marton, Dresselhaus, 2008; 
Dresselhaus et al., 2016; Shin et al., 2021). However, 
the types of fertilization were not considered. 
Fertilization of the central cell by sperm has been 
other structural characteristics are specific. Of considerable scientific interest is the hypothesis of the 
similarity of fertilization processes in plants and 
animals. According to this hypothesis, in plants, 
like animals, there are two types of fertilization, 
depending on when the nuclei of sexual cells unite: 
premitotic (before mitosis of the zygote nucleus) 
and postmitotic (after the onset or during division 
studied in a small number of flowering plants, and 
this process is given less attention than the study 
5


SHAMROV, ANISIMOVA	
v c
m
3
1
g c
2
4
sp
n v c
5
6
7
8
p g
p g
9
10
Fig. 1. Formation of pollen grains and pollen tubes. 1–4 – Ceratophyllum demersum, 5–8 – C. platyacanthum 
(Ceratophyllaceae), 9 – Gagea stipitata (Liliaceae), 10 – Hemerocallis citrina (Hemerocallidaceae). (1–8 – after Shamrov, 
1983; 9 – after Shamrov, 1990a; 10 – after Shamrov, 1990b). g c – generative cell, m – microspore, n v c – nucleus of 
vegetative cell, p g – pollen grain, sp – sperm, v c – vegetative cell. Scale bar, µm: 30.
chromatin and polar nuclei occurs in metaphase, as 
in the postmitotic type (Batygina, 1974; Korobova, 
1982). In Zephyranthes candida Lindl., Z. grandiflora 
Lindl. and Z. macrosiphon Baker (Amaryllidaceae), 
syngamy occurs according to the premitotic type; 
in the first two species the fusion of the central cell 
and sperm is also premitotic, while in Z. macrosiphon 
it is postmitotic (Vorsobina, Solntseva, 1979). In Dios-
corea caucasica Lipsky (Dioscoreaceae) the triple 
fusion is similar to the syngamy of the premitotic 
type. In another species, D. nipponica Makino, an 
intermediate nature of the fusion of sperm and central 
cell was revealed (Torshilova, 2018). 
For a number of flowering plants, it was shown 
that the process of syngamy is simultaneously 
characterized by traits of different types. An electron 
of the egg (syngamy). Since in flowering plants the 
endosperm resulting from triple fusion is necessary 
for coordinating the development of the embryo 
and seed germination, its formation usually begins 
before the division of the zygote. That is why the way 
of endosperm formation does not always correspond 
to the type of fusion of female and male gametes 
during syngamy and most often occurs according to 
the premitotic type or is characterized by signs of 
premitotic and postmitotic types. Thus, for Triticum 
aestivum L. and Zea mays L. (Poaceae), it was shown 
that, although double fertilization occurs according 
to the premitotic type, the sperm chromosomes of 
the resulting nucleus of the primary endosperm cell 
enter prophase separately from the chromosomes 
of the polar nuclei. Complete union of sperm 
	
БОТАНИЧЕСКИЙ ЖУРНАЛ     том 109     № 1     2024


	
GAMETE STRUCTURE AND DEVELOPMENT IN CONNECTION WITH FERTILIZATION...	
7
m
microscopic study revealed that the fusion of the 
sexual nuclei is carried out according to a united plan, 
i
while they actually represent a single lobed nucleus, 
and the patterns of different types of fertilization at 
the light-optical level represent differences in the 
rate of their complete union (Plyushch, 1992).
A comparative analysis of syngamy in plants and 
n
e s
ps
animals revealed that in the postmitotic type in 
plants, sperm enters the egg, while in many animals, 
the spermatozoid penetrates into developing oocyte 
during the first or second division of meiosis and 
activates it for further development (Shamrov, 2015b). 
pd
h
ch
It was the presence of a number of contradictions 
in the literature that prompted us to analyze the 
available data not only on syngamy, but also on 
the features of the formation of sexual cells that are 
involved in its passage. 
Gametogenesis. In flowering plants, male 
v b
gametogenesis is a long process and occurs within 
specialized gametophytes that develop in anthers. 
The male gametophyte is represented by pollen grains, 
2-celled (Fig.  1, 1–4, 9) or 3-celled (Fig. 1, 10). 
In 2-celled pollen grains, gametes are not yet formed, 
and they arise during the growth of the pollen tube 
(Fig.  1, 5–8). In both variants, the gametes are 
Fig. 2. Mature ovule structure in Ceratophyllum demersum 
(Ceratophyllaceae) (after Shamrov, 1997). e s – embryo 
sperm-cells (Shamrov, 2015a). 
Sperms are connected to each other by a common 
sac, h – hypostase, i – integument, m – micropyle, 
n – nucellus, pd – podium, ps – postament, v b – vascular 
bundle. Scale bar, µm: 30.
sac (Dumas et al., 1985). The cytoplasmic projection 
plays a structural role in linking the male germ unit, 
but potentially can perform other important roles 
(McCue et al., 2011). There is an opinion that such 
median cell plate or by protrusions of their protoplasts 
using structures resembling plasmodesmata (Russel, 
Cass, 1981). Detailed studies on sperm formation 
have been carried out. In Hordeum species (Poaceae), 
during the division of the generative cell, organelles 
move to the peripheral regions of the cytoplasm. 
Microtubules, individual cisterns of the granular 
a complex arises already in a 2-celled pollen grain, 
while the nucleus of the vegetative cell contacts the 
cytoplasmic “tail” of the generative cell (a large 
number of microtubules are located here) (Ermakov 
et al., 2016). 
In most flowering plants, the sperms are isomorphic, but in some, their dimorphism is noted – 
the sperms of one pair differ in size, amount of 
cytoplasm, and volume of nuclei. In Plumbago 
zeylanica L. (Plumbaginaceae), the larger sperm 
is in contact with the nucleus of the vegetative 
cell (Russell, 1984). In Nicotiana tabacum, the 
sperm that is not associated with the nucleus of a 
endoplasmic reticulum, and many free ribosomes are 
found in the area of the spindle. In the late telophase, 
vesicles of dictyosomes and microtubules are located 
in the center of the phragmoplast, where the median 
cell plate is formed between the sperms (Charzynska 
et al., 1988). In Nicotiana tabacum L. (Solanaceae), 
the division of the generative cell is also accompanied 
by the formation of a phragmoplast between the 
protoplasts of two sperms (Palevitz, 1993), while 
in Tradescantia virginiana L. (Commelinaceae), 
this process occurs without the participation of the 
phragmoplast and is characterized by the formation 
of a constriction (Palevitz, Cresti, 1989). In flowering 
plants, one sperm cell has a cytoplasmic projection 
with the nucleus of a vegetative cell. The Male Germ 
vegetative cell is smaller, richer in plastids, and 
poor in mitochondria compared to the associated 
sperm (Yu et al., 1992). Dimorphism was found 
in members of other families of flowering plants: 
Unit (MGU) appears and moves towards the embryo 
БОТАНИЧЕСКИЙ ЖУРНАЛ     том 109     № 1     2024


SHAMROV, ANISIMOVA	
a c
1
2
3
t mg
4
5
6
7
e s
8
9
Fig. 3. Ovule structure at archesporium stage (1), megasporogenesis (2–5) and first stages of embryo sac development 
in (6–9) in Gentiana lutea (Gentianaceae) (after Shamrov, 1988). 1–9 – stages of development. a c – archesporial cells, 
e s – embryo sac, t mg – tetrad of megaspores. Scale bar, µm: 30.
Chenopodiaceae – Spinacia oleracea L. (Wilms, 
is associated with male gametes using short actin 
microfilaments (Heslop-Harrisson J., HeslopHarrisson Y., 1989; Russell, 1992; Knox et al., 1993). 
1986), Brassicaceae – Brassica campestris L. and 
B. oleracea L. (Dumas et al., 1985), Poaceae – Zea 
mays (Rusche, Mogensen, 1988), Euphorbiaceae – 
Female gametophytes are represented by embryo 
Euphorbia dulcis L. (Murgai, Wilms, 1988), Ericaceae – Rhododendron macgregoriae F. Muell. (Taylor 
et al., 1989), Liliaceae – Gagea lutea (L.) Ker Gawl. 
sacs of different structures, depending on the type 
of their development. Their formation occurs in the 
ovule (Johri, 1963; Shamrov, 2008; Rudall, 2021). 
(Yang et al., 1995). The contractile proteins (myosin 
The ovule of angiosperms is an organ comprising 
the nucellus, integuments, chalaza, and funiculus. 
The events of archesporium differentiation, megasporogenesis and embryo sac formation take place 
and actin), which are part of microfilaments, were 
found in the cytoplasm of a vegetative cell. Myosin 
is located on the nucleus of a vegetative cell and 
	
БОТАНИЧЕСКИЙ ЖУРНАЛ     том 109     № 1     2024


	
GAMETE STRUCTURE AND DEVELOPMENT IN CONNECTION WITH FERTILIZATION...	
9
show polyploidization of nuclei (Ceratophyllaceae, 
Gentianaceae, Ranunculaceae, etc. – Zhukova, 
Sokolovskaya, 1977; Shamrov, 2008; Butuzova, 
2018) (Fig. 5, 2). In some plants, there is an increase 
in the number of antipodes (Nelumbonaceae – 
Titova, 1988), which is accompanied by an increase 
in it (Fig. 2). After fertilization both the embryo 
and endosperm arise. The complex transformations 
of the embryo, endosperm and surrounding tissues 
developing in conjunction with them result in 
the seed formation. The ovules and seeds are 
characterized by considerable diversity in the 
shape, the degree of development and the structure 
(Shamrov, 2018). Various specific structures are 
in the ploidy of nuclei in cells (Poaceae, etc. – 
Batygina, 1974). Antipodal cells are equipped with 
formed in the nucellus, that provide differentiation. 
The postament is a column-like tissue located below 
complete cell walls. They are organized according 
to the transfer cells, and play an important role in 
the nutrition of the embryo sac, performing the 
functions of adsorption, transport, and synthesis of 
a number of metabolites. 
In the formed embryo sac of many plants, all 
cells of the egg apparatus have a complete cell walls. 
In the process of maturation of the embryo sac in 
the apical parts of the egg and synergids, some of 
the cell walls are lost (possibly, substances cease to 
accumulate in them), and before fertilization, the 
protoplasts of these cells (on the side of the central 
cell) are surrounded only by the plasmalemma 
(Russell, 1992). In the basal part of the synergids, 
the sporogenous or gametophytic structures. The 
podium is a cup-shaped structure arising in the 
chalazal zone of the nucellus. The hypostase is a 
boundary tissue between the nucellus, integuments 
and chalaza. The major function of all the special 
structures in question appears to be that of directed 
translocation of nutrients: the hypostase – to the 
nucellus and integuments; the podium – to the 
lateral (lateral transport through integumentary 
tapetum and central cell) and, presumably, apical 
nucellar regions (apical transport through synergids 
and parietal tissue); and the postament – to the basal 
nucellar region (basal transport through antipodals) 
(Shamrov, 2008, 2022). 
Female gametophyte in angiosperms consists 
a filiform apparatus differentiates – a system of 
winding, highly branched outgrowths of the cell wall 
(Fig. 4, 6; 5, 1). Such outgrowths increase the surface 
of egg apparatus, antipodals, and central cell. The 
egg apparatus is presented by egg and two synergids 
(Female Germ Unit – FGU). This notion was 
proposed by Dumas et al. (1984). Subsequently, 
the content of this concept was expanded. Now 
the female germ unit is comprised of the egg, two 
synergids, and the central cell (Huang, Russell, 
of the plasmalemma, which gives it the properties 
of transfer cells for the transport of substances. The 
central cell in the area of contact with the egg is also 
covered with a plasmalemma. Here, an extracellular 
space, or “gap”, is formed, where sperms enter. It is 
designated as the gamete interaction zone (Sprunck 
et al., 2012).
1992). The female gametes are the egg and the central 
Fertilization traits in angiosperms and 
gymnosperms in the light of endosperm origin. 
In gymnosperms (Goroschankin, 1880) and 
cell (Fig. 3, 1–9; 4, 1–6). The egg is located on the 
side, usually slightly below the synergids. It, as a 
rule, has a pear-shaped shape and is characterized 
by morphological polarity: the nucleus is located 
at the apical end, and the vacuole is located at the 
basal pole. The synergids are egg-like and are also 
characterized by polarity, with the nucleus at the 
basal end and the vacuole at the apical pole or center 
of the cells (Fig. 4, 5, 6; 5, 1–3). In the largest 
central cell of the embryo sac, 2 polar nuclei most 
often form (Fig. 4, 4), which usually fuse to form a 
secondary nucleus before fertilization (Fig. 4, 6; 5, 
1–3). The antipodal cells are arranged in the form 
angiosperms (Strasburger, 1884; Guignard, 1886), 
like other higher plants, single fertilization was first 
described. It was believed that the second sperm of 
the pollen tube is destroyed and does not participate 
in fertilization. However, double fertilization was 
later found in flowering plants, which is one of the 
distinguishing characteristics of these plants. It 
includes the union of one sperm with an egg (later, 
an embryo is formed from the resulting zygote), and 
the second sperm (of the same pair) with the central 
cell of the embryo sac (endosperm develops). 
The phenomenon of double fertilization was 
of a triangle or a line of three cells and are localized 
at the chalazal pole of the embryo sac (Fig. 4, 6; 
5, 1). They may be ephemeral and disappear before 
or during fertilization. Remaining after fertilization, 
the antipodes can increase in size, while they 
discovered in the study of fertilization in Fritillaria 
tenella M. Bieb. and Lilium martagon L. from the 
Liliaceae family (Nawaschin, 1898a, b). Both 
БОТАНИЧЕСКИЙ ЖУРНАЛ     том 109     № 1     2024


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