Polyploids

Distribution and Occurrence of Polyploidy:

▪ Polyploidy is wide spread in plant

kingdom but is less common in animal
kingdom.
▪ Darlington and Janaki-Ammal in 1945 and
Stebbins in 950 estimated that polyploids
make up 30 to 35% of Angiosperms.
▪ high frequency of polyploid specie-
Rosaceae, Polygonaceae, Malvaceae,
Crassuiaceae, Nymphaeaceae;
▪ nearly 75% of the Gramine are polyploid
▪ polyploidy is the most prevalent in
herbaceous perennials
 Polyploidy

▪ The phenomenon of change in chromosome number

is called heteroploidy, which includes aneuploidy and
euploidy.
▪ The change in chromosome number may involve one
or a few chromosomes of a genome, and the
phenomenon is called aneuploidy, or the change may
involve one or more complete sets of chromosomes
or genomes and the phenomenon is called euploidy.
▪ The basic chromosome number of a species is
represented by ‘X” whereas ‘n’ represents the gametic
chromosome number and ‘2n’ represents the somatic
chromosome number of a species.
▪ Euploidy is designated by ‘X’, ‘2X’, ‘3X’, ‘4X’, ‘6X’, etc.
▪ Herteroploid- a change from 2x
A. Aneuploid- One or few chromosomes extra or missing from 2n
▪ Nullisomic 2n-2
▪ Monosomic2n-1
▪ Double monosomic 2n-1-1
▪ Trisomic 2n+1
▪ Double trisomic 2n+1+1
▪ Tetrasomic 2n+2
B.Euploid
▪ Monoploid x
▪ Haploid n
▪ Polyploid More than 2 copies of genome
Autoplyploid—AAA (3x),AAAA (4x),AAAAA (5x),AAAAAA (6x)
AAAAAAAA (8x)
Alloplyploid –AB…Allitetraploid,Allohexaploid &Allooctaploid
Classification of Euploidy
▪ When one or more complete sets of
genomes are involved, the condition is
called euploidy.Euploidy is classified as:
▪ 1.Monoploidy / Haploidy
▪ 2.Diploidy
▪ 3.Polyploidy
▪ • Autopolyploidy
▪ • Allopolyploidy
▪ Autoallopolyploidy
▪ segmental allopolyploidy
▪ Agmatoploidy
▪ 1. a.Autotriploidy (3x)
▪ b. Autotetraploidy (4x)
▪ c. Autopentaploidy (5x)
▪ d. Autohexaploidy (6x)
▪ 2. Allopolyploidy
▪ a. Allotetraploidy (2x1+2x2)
▪ b. Allohexaploidy (2x1+2x2 + 2x3)
▪ c. Allooctaploidy (2x1+2x2 + 2x3 + 2x4)
▪ Autopolypoids:
▪ Polypoids have larger cell size than diploids, larger guard cells of
stomata, larger pollen grains, slower in growth and later in flowering,
larger and thicker leaves, larger flowers & fruits but lesser in
number than of diploids, reduced fertility due to meiotic irregularity
and genotypic imbalances.
▪ Autotriploids are generally highly sterile e.g. water melons, banana
etc. but in some cases they are highly fertile e.g., spinach.
▪ In autotetraploid, 4 chromosomes are homologous to each other
hence each gene has 4 copies.
▪ Autopolypoid played a limited role in the evolution of plant species.
Autopolypoid crops are potato (4x), alfalfa (4x), banana (3x), and
sweet-potato (6x).
▪ Autotriploids and Autotetraploids are found more commonly in
nature than the higher polyploids.
▪ Autotriploids have been reported in grapes, watermelons, sugar
beet, tomato, banana, lemon, apple, etc., where as autotetraploids
have been reported in rye, corn, berseem, marigold, flox,
snapdragon, grapes, apples and several vegetables
 Origin and production of Polyploids
▪ Autopolyploids are generally produced by
chromosome doubling. This chromosome
doubling may occur spontaneously in nature,
or may be induced artificially by physical and
chemical agents.
▪ Spontaneous
▪ Prduction of adventitious buds
▪ Treatment with physical agents
▪ Regeneration in vitro
▪ Colchicine treatment
▪ Other chemical agents
Origin and production of Autopolyploids

Spontaneous production

▪ by the formation of unreduced gametes followed by

their fertilization.
▪ by chromosome doubling followed by failure of cell
division or failure of 2nd meiotic division.
▪ Production of unreduced gametes is promoted by
certain genes causing complete asynapsis or
desynapsis-such mutant genes producing parallel
spindle(ps) and omission of second division (os) in
potato
Formation of a tetraploid organism
 Origin and production of Autopolyploids

Treatment with physical agents

▪ X-ray or gamma ray irradiation
▪ Eg:Datura-cold treatment
▪ Maize-38-45 0c at the time of first division of zygote
produce 2-5% tetraploid progeny
▪ Other eg: barley,wheat, and some crop sps.
▪ Regeneration in vitro
▪ Common in cells cultured in vitro
 Cochicine traetment
▪ From poisonous seed isolated from seeds(.2-
.8%) and bulb(.1-.5%)
▪ It is a chemical compound which was first
discovered by Pernice in 1889. Colchicine was
first demonstrated by Levan (1938) to be a
specific and efficient chemical in creating
polyploid restitution nuclei. Colchicine is obtained
from the extract of seeds (0.2 – 0.8%) and corms
(0.1- 0.5%) of the Colchicum autumnale (member
of family Liliaceae).
▪ In India it is obtained from Colchicum luteum and
Gloriosa superba.
 Effect of Colchicine:

▪ When colchicine is applied some irregularities

occur in the mitotic division. It has no effect
on doubling of chromosomes which proceeds
in normal fashion. This chemical is a spindle
poison and it simply checks the anaphase
movement of chromosomes to the two poles
and consequently mitosis fails to complete.
▪ The doubled chromosomes become bounded
by a nuclear membrane, thus a restitution
nucleus with double chromosome number is
formed.
 Triploids:
▪ Produced by hybridization between tetraploid and diploid strains;
generally highly sterile, useful in production of seedless
watermelons.
▪ Seedless watermelons are produced by crossing tetraploid (4x, )
and diploid (2x, ) lines and are grown commercially in Japan. They
do not produce true seeds.
▪ Triploid sugarbeets have larger roots and produce more sugar per
unit area than diploids. Triploid sugarbeets were grown
commercially in Europe and Japan but their popularity is declining.
Seed production is difficult because beet flower is small.
▪ A triploid (3x) clone of tea (Camelia assamica) has been released for
commercial cultivation in northern India. Triploid cultivar TV 29
produces larger shoots & biomass; yields more cured leaf per unit
area and tolerant to drought than available diploid varieties.
▪ Triploids can’t be maintained except through clonal propagation.
▪ The anaphase segregation of each trivalent is
irregular.
▪ For example the trivalent can segregate in 2:1 or
1:1 (with one chromosome lagging).
▪ The overall chromosome segregation in triploids
is highly irregular, leading to the formation of
defective or sterile gametes, which fail to take
part in fertilization. Thus triploids do not produce
seeds due to their irregular meiotic behaviour.
▪ Triploids can, however, be maintained by
vegetative propagation, or by crossing diploids
and tetraploids.
Problems with
being triploid
failure of
chromosomes
to segregate
properly.
This is why
triploid
watermelons
are seedless.
20
Autotrploi
ds
Fig. 1. Chromosome associations at M I of meiosis in triploid S. tuberosum. (a) 11 III
+ 1 lI + 1 I, (b) 1011I + 2 I I ÷ 2 I , ( c ) 8 I I I + 4 I I + 4 I , and (d ) 41II + 8 I I + 8 I ( a b
Fig. 2. Chromosome distribution at A 1 of meiosis in triploid S. tuberosum. (a)
20-16, (b) 19-16 plus 1 lagging chromosome, (c) 19-15 plus 2 laggards, and (d)
15-15 plus 6 laggards. All lagging chromosomes are dividing precociously (about ×
Fig. 3. Cells at the tetrad stage in triploid S.
tuberosum. (a) normal looking tetrads, (b)
dyads and triads.
Fig. 3. Cells at the tetrad stage in triploid S.
tuberosum. (a) normal looking tetrads, (b)
dyads and triads.
Chromosome squashes of diploid H. nuttallii N102 (A), triploid F1 (2n = 3x =
51) (B), and diploid F 1 (2n = 2x = 34) (C) derived from the cross of nuclear
male-sterile (NMS) HA 89-552/N102. Seedlings (D) of diploid (left) and
triploid F1s (right), and adult plants (E) of diploid (left) and triploid F 1s (right),
flowering capitula of diploid (F) and triploid F1s (G), respectively. Bars = 5
 Tetraploids
▪ autotetraploids may be superior in some
quality characters to their respective
diploids
▪ e.g., tetraploid maize has 43% more
carotenoid pigment and vitamin A.
▪ Certain distant crosses are not successful
at the diploid level, but are relatively
successful at the autotetraploid level
▪ e.g. 4x Brassica oleracea x B. chinensis is
successful.
▪ Autotetraploid varieties of forage crops are
proved successful
▪ In banana tetraploids are inferior than
diploids-weaker leaves and increased
fertility
▪ Tetraploid produced by chance fertilization
of triploid egg AAA and haplid A pollen
▪ Not a commercial success
▪ Maize-43% more carotinoid pigment and
vitamin A
▪ Overcome incompatibility by polyploidy
eg:Tobacco,White clover, Petunia
▪ Distant crosses are not successful at
diploid level,but tetraploid level
▪ Eg:4x B.oleraceae XB chinensis
▪ 2n B.oleraceae is not successful
▪ Larger in size and more vigorous than
diploids
▪ e.g. tetraploid red clover (Trifolium
pratense), rye grass (Lolium perenne),
aliska clover (Trifolium hybridum variety
Tetra) and Berseem (Trifolium
alexandrium) variety Pusa Giant have
been proved most successful.
Autotetraploid
 Tetraploids
▪ Autotetraploid turnips (B. rapa) and cabbage
(B. oleracea) are larger in size and have more
water content than diploids.
▪ Many ornamental plants are autotetraploids
which have increased flower size and longer
flowering duration.
▪ Pusa Giant Berseem is the 1st autopolypoid
variety released for general cultivation in India.
It yields 20-30% more green fodder than
diploid varieties.
 Tetraploids
▪ Sugandha is an autotetraploid variety of
vetiver (Vetiueria zizanoides) which
gives 11% more oil yield.
▪ Variety HMT-1 of Hyoscyamus niger is
an autotetraploid which gives 15% more
biomass and 36% more crude drug yield
than diploid parent.
▪ Seed is the commercial product,4x is less
successful due to high sterility and
genetic instability
▪ Fertility can be improve through the
breeding and selection
▪ Successful for crops with more no. of
chromosomes
▪ Cross-pollinated species are more
responsive
▪ Crops grown for vegetative parts are
successful
▪ Cytology
▪ autotetraploids show tetravalents of various
shapes at metaphase.
▪ Occasionally, there can be two bivalents or a
trivalent plus a univalent at metaphase. At
anaphase, generally there is 2:2 segregation of a
tetravalent producing balanced 2X gametes
which on fertilization result in the production of
tetraploid progeny.
▪ Sometimes, there can be 1:3 or 2:1 (one
chromosome lagging) segregation at anaphase.
▪ , autotetraploids mostly produce fertile gametes;
however, some sterility is also found.
Non functional
gamete

34
). The other combinations are homeologous (
▪ Segrgation of Autotetrapolids

▪ Aaaa-simplex
▪ AAaa-duplex
▪ AAAa-triplex
▪ AAAA-quadruplex
▪ Aaaa-nulliplex

▪ .
 Morphological features and applications of
Autopolyploids
▪ • One of the important features of autopolyploids
is their gigantism. The fruits, flowers, seeds,
leaves etc. are generally larger in size than in the
normal diploids.
▪ larger cell size than diploids. Guard cells of
stomata are larger and number of stomata per
unit area is lower in polyploids than in diploids.
▪ Pollen grains are larger than those of the
corresponding diploids.
▪ generally slower in growth and show delayed
flowering and maturity.
▪ reduced fertility due to irregularities during
meiosis and due to genotypic imbalance leading
to physiological disturbances.
▪ • Different species have different
tolerance levels in ploidy. Some species
survive up to 3X level while others can
tolerate up to tetraploid (4X) and
hexaploid (6X) levels.
▪ • Autotetraploids are produced in those
plants where large size of seeds and
fruits, increased flower and leaf size is
the economic requirement, e.g. several
fruit crops, vegetables, ornamentals and
forage crops.
 Limitations of Autoplyploidy

▪ 1.Larger size is accompanied with high

water content
▪ 2.High sterility accompanied with poor seed
set.
▪ Fertility of autotetraploids can be increased
by hybridization and selection
▪ Triploids cannot be maintained ,except
through clonal propagation,they have to
regularly produced by crossing 4x X 2x
 Limitations of Autoplyploidy
▪ New polyploids have several undesirable
characters
▪ Eg:poor strength in grapes
▪ Irregular shapes of watermelon
▪ So a new polyploids can rarely be used
direcly in crop production
▪ Effect of autopolyploids cannot be predicted.
 Segmental Allopolyploidy
Some degree of homology (partial
homology) may exist between some
chromosome of one genome and those of
the other genome.
Therefore, in such polyploids both bivalents
and multivalents are formed.
This type of chromosome pairing is called
heterogenetic pairing or allosyndetic pairing.
▪
▪ The products may be sterile due to genetic
imbalance. Segmental allopolyploids may show
tetrasomic inheritance for genes on some
chromosomes, while disomic inheritance for
genes on other chromosomes.
▪ Therefore, the disomic and tetrasomic inheritance
in the polyploid are evidences of segmental
allopolyploidy.
▪ Agmatoploidy
▪ The most extensive aneuploid series in the plant
kingdom is that in the genus Carex, in which
haploid numbers ranging from n = 6 to n = 56
have been reported, and every number from 12 to
43 is represented by one or more species

▪ Carex: Cyperaceae exhibit remarkable

agmatoploid chromosome series between and
within species. This chromosomal diversity is due
in large part to the structure of the holocentric
chromosomes:

▪ fragments that would not be heritable in

organisms with monocentric chromosomes have
the potential to produce viable gametes in
organisms with holocentric chromosomes.
 Luzula sect. Luzula in the south-
eastern Alps—karyology and genome
size Tinka Bačič, Nejc Jogan & Jasna
Dolenc Koce TAXON 56 (1) • February
2007: 129–136

Metaphase or late prophase

chromosomes of Luzula.
A, L. campestris (2n = 12 AL);
B, L. divulgatiformis (2n = 24 BL);
C, L. exspectata (2n = 24 BL);
D, L. sudetica (2n = 48 CL);
E, L. divulgata (2n = 24 AL);
F, L. multiflora tetraploid (2n = 24
AL);
G, L. alpina (2n = 12 AL + 24 BL);
H, L. multiflora hexaploid (2n = 36
 B. Allopolyploidy
▪ Origin and production
▪ by hybridization between two or more than two
different species (interspecific or intergeneric)
followed by doubling of chromosomes.
▪ The two or more species can hybridize naturally, or
can also be crossed artificially.
▪ chromosome doubling of the hybrid can occur
spontaneously, or can be induced by colchicine
treatment.
▪ Genomes of two different species cannot survive in
diploid condition in a hybrid due to lack of
chromosome pairing and genetic disharmony. That is
why evolution has favoured chromosome doubling so
that they survive at tetraploid or hexaploid level.
▪ In F1 hybrid due to irregular mitotic
division chromosome number doubled in
certain branches which are fertile and
also allopolyploids
▪ Eg:
▪ Primula verticellata x P.floribunda- certain
branches are allopolyploid and named as
▪ P.kewensis
▪ Brassica oleraceae var.capitata X R.sativa
▪ Experimental production of
allopolyploids can achieved
by chromosome doubling of
the hybrid with the help of
colchicine treatment-termed
as synthetic allopolyploids
 Morphological features of Allopolyploids
▪ Combined the morphological and
physiological features of both the parents
▪ Very dofficult to predict the conbination of
character
Undesirable character conbination
▪ eg;:raphanobrassica-rabbage now radicole
-leaf like radish and root like cabbage
Desirable character conbination
▪ Triticale-hardiness of rye and high yeild of
wheat
▪ In general more vigorous than diploids
▪ Adaptibility differ from parents
▪ May produce some novel secondary
metabolotes
▪ Increased pathogen resistance
▪ May have some novel morphological
characters
▪ Many of them are apomictic eg:Grasses
(Complete sterility of allopolyploids may adapted to apomictic condition)

▪ Two genomes are transferred to the cytoplasm

of one parent.Nuclear-cytoplasmic interaction
play a role in the determination of character
▪ .
▪ Differncial gene expression-diffencially
expressed in different species
▪ . Eg:in Gossypium-out of 40,430 about
53% genes expressed diffentially in AA,BB
&GG genomes
Gene expressed diffencially in parents
have two effects
▪ Additive gene expression-mid-parental value
▪ Non-additive gene expression-deletion if one
parental value-either increase or decrease
▪ One of the two parents act as “expression
dominant” the expression level in the
allopolyploid comparable to the parents
contributed by that parent
▪ Some genes show decreased expression and
some show increased expression in
allopolyploid than parents
▪ Additive effect contributed by
–”expression dominant genome “
▪ Non-additive effect contributed by
“expression recessive genome”
▪ Changes in gene expression may be due
to:
▪ Alteration in the aminoacid sequences of
protein
▪ Differences in gene regulation pattern
▪ Changes in the chromosome structure
▪ Affect the epigenetic changes as DNA
methylation &stable chromatin structure
alteration
▪ Cytological features of allopolyploids
▪ Homoeologous chromosomes
▪ Since allopolyploids (AABB, AABBCC) have two or more
different genomes, they behave like diploids during
meiosis and are thus also called amphidiploids, as the
genomes of two different species behave like diploids.
▪ Sometimes, the genomes of two or more than two
different species have some homology so that
multivalents are occasionally formed during meiosis.
Such polyploids are called segmental allopolyploids.
▪ natural allopolyploids might have been produced due to
following reasons.
▪ • Failure of chromosome segregation at anaphase after
zygote formation.
▪ • Irregular mitotic cell division in apical meristem, leading to
the formation of allopolyploid branch.
▪ • Fertilization between unreduced gametes
▪ Fertility of allopolyploids can be improved
through hybridization and selection.
▪ Increased in the bivalent formation as a
result of selection so that established
polyploids have become identical to
diploids in terms of chromosome
behaviour ie.they have become diplodized.
▪ The process due to which allopolyploids
show diploidlike behaviour ie.only
bivalent formation is known as
‘diplodization’
▪ Role of allopolyploidy in evolution
Synthesis of Triticale & Rephanobrassica:

Hardiness of rye (Secale

cereale) and yielding
ability of wheat (Triticum
turgidum) were combined
in Triticale hexaploide.
But in Raphanobrassica,
the aim was not fulfilled.
The aim in producing
Raphanobrassica was to
synthesize a crop
species that would
combine the root of
radish (R. sativus) with
the leaves of cabbage (B.
oleracea).
Raphanobrassica
combined the leaves of
radish and roots of
cabbage.
Raphanobrassica
 Role of allopolyploidy in evolution
▪ Allopolyploids have been found more
successful as crop species than
autopolyploids due to their better adaptability.
▪ Many of our present day crop species are
allopolyploids, and have thus contributed to a
great extent in the evolution of plants.
▪ It is estimated that about 1/3rd of the present
angiosperms are polyploids, and that majority
of them are allopolyploids.
▪ wheat, brassica, cotton, tobacco, etc. are all
allopolyploids.
▪ Natural allopolyploids are identified by karyotype
analysis and meiotic behaviour.
▪ It has been possible to trace back the evolutionary
history of many allopolyploid crop species and their
diploid progenitors have been identified with some
degree of certainty.
▪ The identification of diploid parents is primarily based
on pairing between the chromosomes of diploid
progenitor and the allopolyploid species.
▪ The homology between some of the chromosomes
suggests that diploid species may be one of the
parental species of the allopolyploid.
▪ Additional evidence on the parental
diploid species is obtained by
synthesising the naturally occuring
allopolyploid from parental diploid
species. The resemblance between
synthetic and natural allopolyploid
▪ . Protein and enzyme analysis and
chromosome banding provide the
further confirmatory evidence.
Evolution of Cotton

Gossypium hirsutum,
also known as upland
cotton or Mexican cotton,
is the most widely
planted species of cotton
in the world. Globally,
about 90% of all cotton
production is of cultivars
derived from this species.
9 old world and 8 new
world cotton
▪ Gossypium barbadense is one of several species of cotton.
It is in the mallow family. It has been cultivated since
antiquity, but has been especially prized since a form with
particularly long fibers was developed
▪ Gossypium arboreum, commonly called tree cotton, is a species of
cotton native to India, Pakistan and other tropical and subtropical
regions of the Old World. There is evidence of its cultivation as long
ago as the Harappan civilization of the Indus Valley for the production
of cotton textiles
▪ Gossypium herbaceum, commonly known as Levant cotton, is a
species of cotton native to the semi-arid regions of sub-Saharan Africa
and Arabia, where it still grows in the wild as a perennial shrub
Evolution of Cotton
▪ Importantly, the new AD offspring's traits were not
a blend of its parents'.
▪ It was different from both parents, and better-
suited to the hot, dry environment along the coast.
▪ Unlike either parent, it could grow in sand, and it
could withstand salt spray and flooding. Its salt-
resistant seeds floated easily. They took root on
islands throughout the Caribbean, and even
Hawaii and the Galapagos
▪ G.hirsutum-AAD1D1
▪ G.barbadenase-AAD2D2
1. Evolution of bread wheat (Triticum
aestivum) Evolution of Bread
wheat (Triticum
aestiuum) Bread
wheat appeared
about 8000 years
ago as a hybrid
between T.
turgidum and T.
tauschii possible
in the regions of
northern Iran and
Armenia, where
natural hybrids
between T.
turgidum & T.
tauschii can be
found in farmer’s
field
Evolution of Brassica(N,U,1935)

71
Evolution of Nicotiana
Thank you