Apomixis

The normal sexual cycle involves two important processes : (i) Meiosis, and (ii) Fertilization. In a sexual cycle a
diploid generation (sporophytic) alternates with a haploid generation (gametophytic). This normal sexual cycle is
called Amphimixis and the plants are known as amphimictic type of plants.
There are another group of plants where normal sexual cycle is replaced by a type of asexual reproduction. The
phenomenon is called apomixis and the plants are called apomictic plants. Winkler defines apomixis as the
substitution of the usual sexual reproduction by a form of reproduction which does not involve meiosis and
syngamy.
For the sake of convenience, apomixis may be divided into four categories.
(a) Non-recurrent apomixis – megaspore mother cell undergoes meiosis and haploid embryo sac is formed.
The new embryo may arise from the
a. Egg (haploid parthenogenesis) without fertilization
b. Some other cell of the gametophyte (haploid apogamy)
Since the plants produced by this method contains a single set of chromosome. The process is not
repeated from one generation to another.
(b) Recurrent apomixis – embryo sac may arise from
a. A cell of the archesporium (generative apospory/diplospory)
b. Some other part of the nucellus (somatic apospory)
There is no reduction in the number of chromosomes (no meiosis) and all the nuclei of the embryo sac are
diploid. The embryo may arise either from –
i) The egg (diploid parthenogenesis) without fertilization
ii) Some other cell of the gametophyte (diploid apogamy)
(c) Adventive embryony/Sporophytic budding – whatever the method by which the embryo sac is formed
(haploid or diploid) embryos do not arise from the cells of the gametophyte but from those of the
nucellus or the integument. No alternation of generation, Diploid tissue of the parent sporophyte directly
give rise to the new embryo. Embryos arise directly from the diploid sporophytic cells (nucellus or
integuments). The sexual embryo sac develops normally, and the zygotic embryo either degenerates or
competes with the apomictic embryo.
(d) Vegetative reproduction – the flowers are replaced by bulbils or other vegetative propagules which
frequently germinate while still on the plant. Plants propagate by a part of their body other than the seed.
Agamospermy – Embryo is formed by some process in which normal meiosis and syngamy have been eliminated.
However, the seed habit is retained as the agent of propagation. There are three types –
(i) Adventive embryony – Gametophytic generation is completely eliminated. Adventive embryos are
distinguishable from the zygotic embryo because of lateral position and absence of suspensor. E.g.
Citrus, Eugenia, Mangifera indica, Euphorbia dulcis. Although pollination and fertilization are not
essential, stimulatory effects of these factors have been found. Endosperm formation is recorded and
may originate by triple fusion or without it.
(ii) Diplosopory – Embryo sac is formed from MMC without undergoing meiotic division. It may be
divided into two types –
a. MMC undergoes a division which is semi heterotypic. There is no pairing or some degree of
pairing. Restitutional nucleis is formed. The nucleus remains diploid. Ixera, Taraxacum. The
nucleus divides by three mitotic divisions to form eight-nucleate embryo sac.
b. MMC divides directly by mitotic divisions. E.g. Eupatorium, Calamogrostis.
(iii) Apospory – Somatic cell in the nucellus directly forms an unreduced embryo sac, and the diploid egg
parthenogenetically develops into embryo. E.g. Hieracium, Parthenium, Rubus, grasses.
Vegetative reproduction – some form of vegetative reproduction is found associated with the seminiferous plants
(reproducing by means of normal seeds). The plants can be regarded apomictic when vegetative reproduction
replaced the sexual methods. Vegetative reproduction takes place by means of propagules such as bulbils, bulbs,
runners, suckers and so on. There are three types of vegetative reproduction (Gustaffson) –
(a) Propagules formed outside the floral regions and despite the occurrence of functioning sex organs no
fertilization or seed setting takes place. Agave Americana, Elodea Canadensis
(b) Propagules are formed outside thefloral region and the plants are sexually sterile. Fritillaria, Lillium
bulbiferum

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 (c) Propagules are formed on the floral branches either in addition to the flowers or in place of them. The
phenomenon is commonly described as vivipary (vegetative vivipary to distinguish them from mangrove
plants) e.g. Deschampsia, Festuca, Poa, Allium.
Parthenogenesis – Formation of embryo from an unfertilized egg is called parthenogenesis. In autonomous
apomicts, the development of embryo is independent of the pollination stimulus. In some plants, embryo develops
only after the pollination. Then the phenomenon si called pseudogamy. E.g. grasses. Three probable role of
pollination in pseudogamy are –
I) It may stimulate the growth of ovule and ovary
II) It may supply one male nucleus for the development of endosperm, and
III) It may stimulate parthenogenesis
Causes of apomixis - most of the obligae apomicts are of hybrid origin and also polyploidy. Hybrid origin is
indicated during meiosis which is greatly disturbed and accompanied by the failure of synapsis and normal
contraction of chromosomes, In Allium carinatum, the replacement of flowers by buibils is due to a single gene
(recessive gene).
In Parthenium, diploid individuals are sexually reproducing but polypllids are obligate apomicts. Three pairs of
gene determine the breeding behavior. Gene a leads to the formation of unreduced egg, gene b prevents
fertilization and gene c causes the egg to develop without fertilization. Plants with aaBBCC form unreduced eggs
but cannot develop into embryos without fertilization. Plants with genotype AAbbCC produc reduced eggs but
embryos are not formed because fertilization is prevented. Plant with AABBcc genotype have normal sexual
behavior. Only plants with aabbcc will be apomictic.
In facultative condition, environmental conditions play important role.
Significance of apomixis
Apomixis offers the possibility of indefinite multiplication of specially favourable biotype without any variation due
to segregation or recombination. In obligate apomicts the advantage is at the cost of long term evolutionary
flexibility of sexual reproduction. In facultative apomicts or group of plants where sexual and apomictis members
co-exist, the phenomenon is important.

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