Lecture-4

Polyembryony, apomixis, chimeras and bud- sports

Learning objective
 To get acquainted with polyembryony, apomixis, chimeras and bud- sports
 Horticultural significance of polyembryony, apomixis, chimeras and bud- sports

Polyembryony
Polyembryony means that more than one embryo develops within a single seed. It is
also known as adventitious embryony (Nucellar embryony or Nucellar budding).
 Polyembryony can develop from several distinct causes. Specific cells in the nucellus
or sometimes with integument have embryos. Genetically, these embryos have the
same genotype as the parental plant and are apomictic.
 Adventitious embryony occurs in many plant species but is most common in citrus
and mango. In these species, both zygotic and apomictic embryos are produced. In
other species (e.g. Opuntia), no pollination or fertilization is needed.
 Polyembryony is common in mango and citrus. In trifoliate orange (Poincirus
trifoliata) several seedlings arise from one seed.
 Of these seedlings, one seedling, usually the weakest may be sexual, and the others
arise apomictically from cells in the nucellus, which are diploid copies of the mother
plant.
Horticultural significance of polyembryony
Nucellar seedlings in citrus are completely free from viruses, because the embryo sac
and adjoining tissues are impregnated at flowering time with some unknown powerful
substances which kills all the viruses. For immediate requirement of planting material,
development of nucellar lines is the quickest and easiest method. The major possible
horticultural applications of polyembryony are:
 Nucellar seedlings are true-to-type seedlings
 Such seedlings are genetically uniform and can be used as virus free
rootstocks
 More vigorous seedlings – continuous vegetative propagation leads to decline
in vigour in citrus
 Development of virus free seedlings and bud wood
 Significance in breeding programme

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Apomixis
In some species of plants, an embryo develops from the diploid cells of the seed and
not as a result of fertilization between ovule and pollen. This type of reproduction is known
as apomixis and the seedlings produced in this manner are known as apomicts.
 Apomictic seedlings are identical to mother plant and similar to plants raised by other
vegetative means, because such plants have the same genetic make-up as that of the
mother plant.
 Such seedlings are completely free from viruses. Plants that produces only apomictic
embryo and are known as obligate apomicts and those which produce both
apomictic and sexual seedlings are called facultative apomicts.

Types of Apomixis: Maheshwari (1950) classified apomixis into four groups:

1. Recurrent Apomixis: In this type of apomixis, the embryo develops from the diploid
egg cell or from the diploid cells of the embryo sac without fertilization. As a result,
the egg has normal diploid number of chromosomes, just like the mother plant
(Fig.4.1).The species, where recurrent apomixis commonly occur are, Parthenium,
Rubus, Malus, Allium , Rudbeckia, Poa, Taraxacum ,etc.
2. Non-Recurrent Apomixis: In this case, the embryo develops either from the
haploid egg cell or from some other haploid cells of the embryo sac. In this case,
haploid plants are produced, which contain only one set of chromosome of the
mother plant. Hence, the haploid plants are sterile in nature and cannot be
normally perpetuated into the next generation. Non-recurrent apomixis occurs
only in a few species such as Solanum nigrum, Lilium spp. etc.
3. Nucellar Embryony or Adventitious Embryony: In this type of apomixes, the
embryos arise from diploid sporophytic cells outside the embryo sac i.e. cells of
the nucellus, integuments etc. This type of apomixis is quite common in citrus and
certain varieties of mango, where fertilization occurs normally and sexual plus a
number of apomictic (nucellar) embryos develop.
4. Vegetative apomixis or bulbils: In some species of plants, such as Allium ,
Agave, Poa etc., the flowers in an inflorescence are replaced by bulbils or
vegetative buds, which sprout, while still on the mother plant and turn into new
daughter plants.

Advantages
 Assured reproduction in the absence of pollinators, such as in extreme environments
 Maternal energy not wasted in unfit offspring (cost of meiosis)

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  Some apomictic plants (but not all) avoid the male energy cost of producing pollen

Disadvantages
 Can't control accumulation of deleterious genetic mutations
 Usually restricted to narrow ecological niches Lack ability to adapt to changing
environments

Clone
The term clone may be defined as a group of genetically uniform individuals, derived
originally from a sexually produced individual or from mutations and maintained exclusively
by asexual means from one ancestor.
 The common examples are Bartlett pear, Delicious apples, Sultana grapes and
Dashehari mango.
 The goal of vegetative propagation is to reproduce progeny plants identical in
genotype to a single plant. The biological process is known as cloning and the
resulting population of plants is called a clone.
 The uniformity of individual plants, within a clonal population is a major advantage
of clonal cultivars of fruit and nut crops.
 Clonal propagation helps in fixing genotypes, uniformity of population, facilitates
propagation and reduces juvenile phase, combine more than one genotype into single
plant (grafting) and control phases of development.
 If environmental conditions are favourable and the clone is managed properly, its
trueness-to-type can be maintained for hundreds of years.
Genetic variations in a clone
Mutation
Mutation is a single-step genetic change or sudden heritable change within cells of a
clone. In general, these changes take place spontaneously, in the plants regular but rare
intervals.
 A mutation is a genetic change involving some part of the DNA molecule. Genetic
mutations result from structural changes in the nuclear DNA of the chromosome in
the nucleus. DNA also occurs in mitochondria and chloroplasts and defects here can
produce genetic changes.
 Chromosomal changes may be due to chance rearrangement of the four bases in the
DNA molecule (point mutation) rearrangement of different parts of the chromosomes
(deletion, duplication, translocations and inversions), addition or subtraction of

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 individual chromosome (aneuploidy), or the multiplication of entire set of
chromosome (polyploidy).
 The rate of mutation can be increased by treatment with specific mutagenic agents
e.g. X-ray, gamma rays and certain chemicals.
Bud- sports or bud mutations
When mutation occur and suddenly appear as a chance in the branch of a plant, is
called bud sport or bud mutations, because they appear to have originated within a single
bud.
 Detection of a new mutant within a clone may require a series of vegetative
propagated generations and multiple propagations from many buds of the same plant.
 Many ‘sports’ have become commercial cultivars. Mutations may affect fruit (colour,
shape, time of maturity), tree structure (spur type), time of bloom, and as a host of
other traits. Sometimes these mutants have highly useful horticultural traits and have
given rise to important new cultivars (‘Ruby Red’ grapefruit, red coloured sports of
apple and pear.)
 On the other hand, mutations may be undesirable and give rise to misshapen fruits,
low production and susceptibility to diseases.

Chimeras
When a mutation occurs within a single cell of a clone, it initially produces an ‘island
‘of mutant cells within a growing point of a stem. The plant becomes a, mixture of two
different genotypes. This structural arrangement is known as chimeras.
 It is the most important kind of genetic variant within clones typified by various kinds
of variation. The name chimera was historically given to certain unique clonal
variants now known to have arisen as graft chimeras.
 Chimeras develop because of the unique architecture of the apical meristem and the
strategic location of the mutation in a dividing cell near the apex of the apical
meristem.
 The three important types of chimeras are based upon the distribution patterns of
mutated and non-mutated cells.
 Buds arising at different positions on a sectorial chimera may produce shoots
consisting of mutated and non-mutated cells, entirely of mutated cells or entirely of
non- mutated cells or sectorial (rarely), mericlinal or periclinal chimeras as described
herewith.

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 Periclinal chimera: The mutated tissues occupy layers of cells that completely
surround an inner core of non mutated tissue. For examples in many red colored fruit
cultivars in apple (Plate 4.2), the red pigment is located only in the epidermal layers;
whereas the cells of the inner tissue have alleles for green or yellow colour. Similarly,
some black berry (Rubus spp.) cultivars are thorn less; because the cells making up
the epidermis do not have this allele. Periclinal chimeras are relatively stable if
propagated stem or grafting.
 Mericlinal chimera: This combination is similar to the periclinal expect that the
cells carrying the mutant gene occupy only a part of the outer cell layer. In case of
a red mutant on a yellow delicious fruit, the surface of the fruit may have
longitudinal streaks or sectors of red on an otherwise yellow surface (Plate 4.3).
This type is unstable and tends to change into periclinal chimeras, revert to the
non-mutated form, or continue to produce mericlinal shoots.
 Sectorial chimera: The mutated cells in this combination occupy an entire sector
of the stem including all layers of the shoot apex. Sectorial chimeras appear if the
mutation occurs in roots and very early stages of embryos where the cells of the
growing point do not occur in layers. In general, this type is unstable and tends to
revert to mericlinal and periclinal chimeras.
 Graft Chimera: In horticulture, a graft-chimera may arise in grafting at the
point of contact between rootstock and scion and will have properties intermediate
to those of its parents. A graft-chimera is not a true hybrid but a mixture of cells,
each with the genotype of one of its parents is a chimera. Hence, the once widely
used term "graft-hybrid" is not descriptive; it is now frowned upon. In practice
graft-chimaeras are not noted for their stability and may easily revert back to one
of the parents.

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