Apomixis

Introductory article
Article Contents

Abed Chaudhury, Commonwealth Scientific and Research Organization (CSIRO) Plant Industry,
Canberra, Australia

. Introduction

Anna Koltunow, CSIRO Horticulture Unit, Glen Osmond, Australia

ES Dennis, CSIRO Plant Industry, Canberra, Australia
WJ Peacock, CSIRO Plant Industry, Canberra, Australia

. Types of Apomixis
. Importance of Apomixis to Agriculture

Apomixis is the formation of seeds by asexual processes. Apomixis comprises an ensemble

of developmental processes that together alter the reproductive functions in the ovule of
flowering plants, converting the sexual programme to an asexual one. The result of these
alterations is that the apomictically derived embryo is not a product of male and female
gamete fusion as occurs in sexual reproduction but is derived solely from cells in maternal
ovule tissues.

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.

Importance of Apomixis to Agriculture

Figure 1 Apomictic reproduction compared to sexual reproduction in

flowering plants. In sexual reproduction the megaspore mother cell (mmc)
undergoes meiosis, spore selection (ss), and mitosis to produce the
common eight-nucleate polygonum-type embryo sac (esc). The embryo
and endosperm compartments of the seed initiate development after
double fertilization (df) and give rise to mature seed (ms). In the panel
showing diplospory, the Antennaria type is depicted where the mmc
undergoes mitosis to form the embryo sac and endosperm and embryo
production are autonomous. In other diplosporous plants fertilization of
the polar nuclei (pseudogamy) is necessary for endosperm production. In
apospory, the initials (ai) differentiate near the mmc, which may or may not
have undergone meiosis. Sexual and aposporous embryo sacs may coexist
or the sexual embryo sac may degenerate during formation. The
aposporous embryo sac may degenerate during formation. The
aposporous embryo sac can differ in structure to that in sexual species, as
shown by the four-nucleate Panicum-type (p) as illustrated. Seed formation
may be autonomous or pseudogamous. Polyembryony (pe) may be
evident if multiple embryo sacs form. In adventitious embryony, numerous
adventitious initial cells (avi) differentiate next to an embryo sac derived by
sexual processes. After double fertilization of the sexual embryo sac the
numerous developing adventitious embryos gain access to the nutritive
endosperm and seed contains multiple embryos.

ous initial cells dierentiate they immediately enter mitosis

to produce an embryo sac. Some ovules can contain
multiple embryo sacs and, depending on the species, the
structure of the embryo sac may be quite dierent from
that seen in the sexual process. The initiation of the
aposporous embryo sac can occur together with a sexual
one or it can displace or inhibit sexual embryo sac
formation. Termination of the sexual process is most
complete if the aposporous initial cells dierentiate early,
around the time of mmc meiosis, producing an unreduced
apomictic embryo sac.

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

Apomixis is of great importance to agriculture primarily

because of its potential to facilitate multiplication of F1
hybrid seeds. Hybrid seeds are preferred for use in modern
agriculture mainly because they provide a yield advantage.
Hybrid seed is usually produced from two inbred parental
lines and outperforms the yield and vigour of either of the
parental lines. Currently production of hybrid seed is
expensive because the parental lines need to be maintained
as pure lines and during normal sexual development the
hybrid vigour present in the F1 seed breaks down in
subsequent generations because of assortment and recombination, thus necessitating purchase of hybrid seed each
season. If apomixis were to be available as a controlled tool
where it could be switched on and o as required in plant
breeding, hybrid vigour could be xed, enabling indenite
multiplication of the hybrid of uniform quality without
decrease in the yield advantage.
Controlled apomixis would enable the use of a greater
range of parent plants in breeding plants for specialized
agricultural habitats. The current world use of hybrids
such as maize (corn) and rice is based on and relatively
restricted by a rather small base of inbred parental lines.
Furthermore, these hybrids do not perform well under all
conditions. Use of apomixis following crosses between two
candidate parental lines would generate progeny that
would be immediately xed in a hybrid genotype and could
be trialled for productivity. In this way apomixis could
eliminate dependency on current inbred lines and enable us
to expand our current agricultural genetic base.
Apomixis would also remove the need for pollination for
fruit and seed production. Adverse environmental conditions such as drought and cold inhibit pollen formation
and result in crop losses in sexually reproducing systems; in
apomictically grown plants these losses could be avoided.
A signicant proportion of the assimilate of the growing
plant is channelled into the production of the male
reproductive apparatus and gametes; these could be
redirected towards increasing yield. Apomixis will also
facilitate uniform clonal seed propagation of many plants
that are currently vegetatively propagated, including fruit
trees, plantation timbers, and potatoes.
Due to these and other potential benets of apomixis in
agriculture, apomixis is likely to be an important breeding
tool for the future. Little is known about the genes

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net

Apomixis

controlling apomixis. Research is currently being pursued

to isolate these genes and to understand how the process of
apomixis can be regulated to result in high yields of clonal
seed. The value of apomixis will only be realized if it can be
eciently controlled under conditions of commercial
production.

Further Reading
Asker SE and Jerling L (1992) Apomixis in Plants. Boca Raton, FL: CRC
Press.

Chaudhury AM, Craig S, Dennis ES and Peacock WJ (1998) Ovule and

embryo development, apomixis and fertilization. Current Opinions in
Plant Biology 1: 2631.
Koltunow AM (1993) Apomixis: embryo sacs and embryos formed
without meiosis or fertilization in ovules. Plant Cell 5: 14251437.
Koltunow AM, Bicknell RA and Chaudhury AM (1995) Apomixis:
molecular strategies for the generation of genetically identical seeds
without fertilization. Plant Physiology 108: 13451352.
Koltunow AM, Soltys K, Nito M and McClure S (1995) Anther, ovule,
seed and nucellar embryo development in Citrus sinensis cv Valencia.
Canadian Journal of Botany 73: 15671582.
Nogler GA (1984) Gametophytic apomixis. In: Johri BM (ed.)
Embryology of Angiosperms, pp. 475518. Berlin: Springer Verlag.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net