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Introductory article
Article Contents
Abed Chaudhury, Commonwealth Scientific and Research Organization (CSIRO) Plant Industry,
Canberra, Australia
. Introduction
. Types of Apomixis
. Importance of Apomixis to Agriculture
Introduction
Apomixis can best be described in reference to the sexual
process. During sexual reproduction, developmental steps
occurring inside the ovule produce the female gametophyte
(embryo sac) and following a double fertilization event give
rise to embryo and endosperm structures. Steps in the
formation of the most common type of embryo sac
(polygonum-type) include the dierentiation of the megaspore mother cell (mmc) from a vegetative nucellar cell,
meiosis to convert the mmc into four spores, retention of
one of these spores and the degradation of the other three.
Finally, the elaboration of an embryo sac occurs after three
mitotic divisions, nuclear migration and cellularization
events to produce a seven-celled, embryo sac, containing
eight nuclei. When the two sperm cells enter the embryo sac
through the micropylar end, they selectively fuse with the
egg and central cell nuclei to produce the embryo and
endosperm, respectively. The embryo and endosperm
compartments dier in the ratio of their maternal and
paternal genomic content. In the embryo the ratio is one
paternal genome to one maternal genome, while in the
endosperm the ratio is one paternal to two maternal.
Apomixis modies the processes described above and
produces a functional female gametophytic structure that
precludes the sexual assortment of genes and recombination of genes associated with meiosis. In apomixis the
double fertilization event does not occur and the embryo
develops autonomously from the unreduced female
gamete. In some apomicts the endosperm develops
autonomously while in others (pseudogamous apomicts)
fertilization of the central cell by a sperm cell may be
required to produce a functional endosperm.
Types of Apomixis
Three types of apomixis are generally recognized
diplospory, apospory and adventitious embryony. These
apomictic processes are depicted compared to sexual
processes in the formation of a common polygonum-type
embryo sac in Figure 1.
Diplospory
In diplospory, the unreduced embryo sac is derived from
the megaspore mother cell either directly by mitotic
division or by aborted meiotic events. Three major types
of diplospory have been reported, named after the plants in
which they occur, and they are the Taraxacum, Ixeris and
Antennaria types. In the Taraxacum type, meiotic prophase
is initiated but then the process is aborted resulting in two
unreduced dyads one of which gives rise to the embryo sac
by mitotic division. In the Ixeris type, two further mitotic
divisions of the nuclei to give rise to an eight-nucleate
embryo sac follow equational division following meiotic
prophase. The Taraxacum and Ixeris types are known as
meiotic diplospory because they involve modications of
meiosis. By contrast, in the Antennaria type, referred to as
mitotic diplospory, the mmc does not initiate meiosis and
directly divides three times to produce the unreduced
embryo sac.
Apospory
In apospory, the nucellar cells that give rise to the
apomictic embryo sac, termed aposporous initials, are
distinct from the mmc. They are similar in appearance to
the mmc and may dierentiate close to the mmc and
develop into an apomictic embryo sac. Once the apospor-
ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net
Apomixis
the adjoining sexual embryo sac and subsequent endosperm formation is necessary to form viable seeds. The
developing embryos closest to the embryo sac grow
towards it, presumably to obtain nutrient and other
developmental signals from the embryo sac. Adventitious
embryo formation is rapid; often multiple embryo form
and these can hinder development of the zygotic embryo.
Adventitious embryony
In this process embryos initiate parthenogenetically outside of an embryo sac structure. Adventitious embryony is
most commonly initiated later in ovule development from
nucellar and integument tissues. In general, fertilization in
2
ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net
Apomixis
Further Reading
Asker SE and Jerling L (1992) Apomixis in Plants. Boca Raton, FL: CRC
Press.
ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net