ISSN 1021-4437, Russian Journal of Plant Physiology, 2010, Vol. 57, No. 5, pp. 720–731. © Pleiades Publishing, Ltd., 2010.
Original Russian Text © N.A. Moiseeva, V.N. Serebryakova, L. Nardi, S. Lucretti, A.M. Nosov, 2010, published in Fiziologiya Rastenii, 2010, Vol. 57, No.5, pp. 771–782.
720
INTRODUCTION
Investigations of somatic embryogenesis in vitro
aim at producing high-quality somatic embryos capa-
ble to properly complete their development. Although
in vitro somatic embryogenesis in the representatives
of the genus Citrus was described long ago, conditions
that would ensure mass regeneration of appropriate
somatic embryos have not been worked out [1–4]. In
many respects, this depends on the fact that realiza-
tion of the program of embryonic development in vitro
is not synchronous; therefore, the population of
somatic embryos thus produced is permanently heter-
ogeneous. Such a situation is common in polyembry-
onic seeds of citrus plants where numerous somatic
embryos develop from the initial cells of nucellus along
with sexual embryo [5, 6]. It is generally accepted that
because of competition for space and nutrition, devel-
opment of the majority of embryos is suppressed at
early stages and only some of them become differenti-
ated. The mature polyembryonic seeds (MPS) of sweet
orange comprise 2 to 9 embryos of various size, with a
sexual embryo may be present among them. The size
of both nucellar and sexual embryos occurring in MPS
of citrus plants depends on the extent of development
of their cotyledons and ranges from 2 to 9 mm [7]. A
negative correlation was shown between the number
and size of the embryos present in one seed: the more
numerous are the embryos, the smaller they become
RESEARCH
PAPERS
Morphogenetic Status of Somatic Embryos
of Citrus sinensis from Mature Polyembryonic Seeds
and Those Produced In Vitro
N. A. Moiseeva
a
, V. N. Serebryakova
a
, L. Nardi
b
, S. Lucretti
b
, and A. M. Nosov
a
a
Timiryazev Institute of Plant Physiology, Russian Academy of Sciences,
Botanicheskaya ul. 35, Moscow, 127276 Russia;
fax: 7 (495) 977-8018; e-mail: morphogenesis@mail.ru
b
Italian National Agency for New Technologies, Energy and the Environment, Rome, Italy
Received March 3, 2009
Abstract—In tissue culture of sweet orange (Citrus sinensis (L.) Osbeck, cv. Tarocco), we obtained mass
regeneration of somatic embryos with two morphologically distinct cotyledons about 3 mm in length, their
numbers amounting to 110–150 embryos per petri dish and 60 to 80% of the population. The morphogenetic
state of somatic embryos was compared using the embryos with the cotyledons of different size (from 3 to
10 mm) from mature polyembryonic seeds as a test system and the cell number, size, and ultrastructural orga-
nization, and the number of protein bodies in the cotyledon cells as morphological and biochemical criteria.
Cell number in the cotyledons of different size was related to the content of protein bodies therein. Typical
protein bodies where 33 kD polypeptide of storage proteins was identified were detected only in the cotyle-
dons, which size was identical to that of embryonic cotyledons from monoembryonic seeds of citrus plants.
In the cells of smaller cotyledons, we detected protein-accumulating vacuoles with electron-dense inclusions
that irrespective of their size, shape and structure accumulated the gold label. The number of the cells with
protein depositions in vacuoles decreased when the cotyledons became smaller. Irrespective of the origin of
embryos (in vivo or in vitro), lipids were the major storage metabolites in the cells of 3-mm cotyledons. As the
cotyledon-forming cells became smaller and less numerous, their metabolic activity tended to decrease in
line with the fragmentation of endoplasmic reticulum, the absence of polysomic complexes, and indistinct
inner organization of mitochondria and plastids. It was concluded that somatic embryos developing in vivo
and in vitro were physiological dwarfs, that is, the structures with diminutive storage organ with characteris-
tically incomplete cell differentiation. Apparently such forms emerged due to the suppression of cotyledon
growth at the initial stages of their organogenesis; as a result, the cell population could not properly realize
both organo- and histogenesis.
Key words: Citrus sinensis, somatic embryogenesis in vitro, polyembryonic seeds, cotyledons, cell number, cell
size, ultrastructure, protein bodies.
DOI: 10.1134/S102144371005016X
Abbreviations: EC1—embryos of class 1 from mature polyembry-
onic seeds; EC2—embryos of class 2 from mature polyembryonic
seeds; EC3—embryos of class 3 from mature polyembryonic
seeds; MPS—mature polyembryonic seeds; SE-in vitro—
somatic embryos produced in vitro.