Acknowledgement

I would like to thank all the people who helped and supported me with writing this
dissertation project.
Firstly, I would like to express my deepest gratitude to my Supervisor, Dr. R.N. Singh
for his unwavering support, collegiality and mentorship throughout this project. Without
his guidance and persistent help this dissertation would have not been possible.
I would also like to thank our college librarian who provided me the required books
which helped me a lot to prepare my dissertation project.

Contents
1. Introduction
2. Aneuploids or Heteroploids;
Monosomic
Nullisomic
Trisomic
Tetrasomic
3. Euploidy
Haploidy/Monoploidy
Diploidy
Polyploidy
3. Types of polyploids a. Autopolyploids
b. Allopolypliods
5. Application of Polyploids.
6. Role of polyploidy in plants
7. Conclusion
8 References.

INTRODUCTION
An individual carrying chromosome numbers other than true monoploid or diploid
numbers is called heteroploids (Sharp, 1934). Heteroploidy is divided into euploidy and
aneuploidy.
In polyploids, x is the basic (monoploid) chromosome number, n is the gametic
chromosome number of chromosomes, and 2n is the Zygotic or somatic chromosome
number for Ex- the genomic formula of Triticum aestivum is 2n=2x=14. In both cases,
the basic chromosome number x=is seven.
The basic set of chromosomes in a diploid is called a genome.
Every cell in the body of living organism is provided with a packet of genetic material
(the nuscles) which contains chromosomes. The number of chromosomes in the nulei of
a given species is fixed.
The following chart gives the idea about the chromosome number in different organisms.
species of plants and animals have identical chromosome numbers. In such cases one
should not think that these organism will show similar characters, because it is not the
number of chromosomes which differentiates various species from one another, but
rather them nature of hereditary material in the chromosomes determine the character of
species. In most of plant and animals the chromosome number ranges between 1O and
5o.The number above and below this range are comparatively rare. The highest number
of chromosomes reported so far is about 1OO found in Radiaolaria , an unicellular
marine protozoa. In plants, chromosome number ranges from two pairs (in a member of
compositeae, Haplopappus gracilis) to several hundreds found in ferns. In ophioglossum
3

768 chromosomes have been counted in each nucleus. In 14 chromosomes of pea there
are 7 chromosomes of different shapes, sizes and structures and each kind of
chromosome has its own homologue. So there are 7 pairs of chromosomes in pea. This
basic number is denoted by the letter x or n. the individuals in which the somatic cells
have chromosomes in pairs are called diploid (2n). when the Reduction division takes
place in a diploid cell, the daughter nuclei so formed will have chromosome numbers
just half the diploid number, i.e. each daughter cell now gets one member of each
chromosome pair. When two gametes, each carrying haploid number of chromosomes,
fuse they produce a diploid cell. The diploid cell produces the diploid body. In this way
in the life cycle of an individual haploid and diploid conditions come regularly one after
another and the same chromosomes number is maintained from generation to generation.
Several cases, however, iare known in which chromosome number become changed in
the somatic cell this condition is known as ploidy.
(A) Origin and artificial production of autopolyploidsThe autopolyploids may accur in nature or may be artificially produced. When they
are found in nature, their autopolyploid nature is inferred mainly by their meiotic
behavior. Ex. Natural autopolyploidy relevant to northern India, pertains to common
doob grass (cynodon dactylon). In U.P. and Bihar, common doob grass was found to
be an autotriploids. It is perhaps successful due to efficient vegetative reproduction,
because, as will be seen autotriploids are normally sterile and set no seeds.
(B) Effects of chromosome doubling:One of the important effects of polyploidy is to produce gigantism.The
autopolyploids may be larger in size. Sometimes the plants may be smaller than diploids,
but the leaves and flowers and the cells themselves may be bigger in size. Some
important effects are as follows:i)
With the increase in cell size. The water content increases leading to decrease
in osmotic pressure. This results into loss of resistance against frost etc.

ii)

The growth rate decrease due to slower rate of cell division. This leads to a

iii)
iv)

decrease in auxin supply and decrease in respiration.

The time of blooming is delayed and prolonged due to slow growth rate.
At higher ploidy level- autoocotaploids or higher the adverse effects are highly
pronounced and lead to the death of the plants.

Induction of Polyploidy:In recent years, a number of methods have been worked out to induce polyploidy in
plants. Some of them are described below.
1. By Radiation:-

Polyploidy can be induced in plants by exposing their certain parts, such as vegetative
buds and flower buds, to ultraviolet rays, X-rays or other rays of shorter wavelengths.
Irradiation increases the rate of cell division and also cause the multiplication of
chromosome number.
2. By Injury:When the merismatic zones of a plant are injured the cells at the points of injury grow
rapidly and form a callus. Callus growth is enhanced by a chemical substance named
coumerine which also brings about somatic doubling of chromosomes. Vegetative buds
generally developing from callus tissus are ployploid in nature, from injured part of
tomato plants it is possible to produce tetra ploids plants.
3. By Chemical Treatment:A number of chemicals are now known which induce polyploidy in plants. Important
among them are colchicine, granosan. Chloroform, chloral hydrate, some narcotics and
alkaloids, Acenaphthene. Indolacetic acid, Ethyl-mercury chloride etc. colchicine is the
best chemical for this purpose. It is a chemical compound which was first discovered by
Pernice in 1889.
Colchicine was first demonstrated by levan to be a specific and efficient chemical in
creating polyploidy restitution nuclei. Colchicine is obtained from the extract of roots
and corms of the colchicine outumnale. In india. It is obtained from colchicium luteum
and glorios a superba.
Methods tested for colchicine treatment
1. Twigs could be immersed in solutions of various concentration for difference periods
2. Seeds were immersed in colchicine solution for different periods

3. Seeds were germinated on blotting paper soaked in different concentrations of

colchicine solution.
4. Young flower buds could be immersed in test tubes filled with different
concentrations of colchicine for different periods.
5. Vegetative buds could be immersed in 0.5 per cent and 0.25 per cent aqueous
solutions of colchicine for different periods
6. Colchicine could be applied as a paste in anhydrous lanolin with 0.4-0.5 per cent
solution.
7. Colchicine solution could be sprayed on the growing buds.
8. Growing buds could be wetted with aqueous solution.
9. Solution of colchicine in agar may be applied to growing buds on interval.
Adaptability of plants to colchicine treatment
The different crop plants show different range of adaptability to the induction of
polyploidy by colchicine treatment. Not the least, even different genotypes within a
species too show different response to double the chromosomes. According to loue
(1953) and Levan (1948).
1. Plants low in chromosome numbers are more likely to respond well in doubling the
chromosome than those higher in chromosome number.
2. Cross-fertilizing plants are more likely to successful for doubling the chromosome
numbers than self-fertilizering species.
3. Vegetatively propagated plants may be more successful as autopolyploid than seed
grown plants.
4. Fertility of autotetraploids may be reduced to a great extent.

Effects of treatment
The changes brought by chromosome doubling are of great significance. Both
individually as well as doubling of chromosomes bring morphological, physiological
and genetic changes.
Morphological : Leaves become large, darker in colour, leathery in texture : leaf lobes
get overlapped and twisted. Stomata, bracts, Pollen grains, bolls and seeds become
bigger than control. Maturity is delayed.
Immediate effect of seed treatment causes swelling of radicles and retardations of their
growth. Seedling show slow growth with thicker stem, darker and broader leaves.
Size of pollen grain and guard cells are bigger than control plants.
The stems of polyploids are stouter and thicker, leaves are larger and broader, hair on the
vegetative parts are coarse and thicker. The floral, fruits and seeds are larger than
diploid.
Normally, polyploids form gigas character because individual cell size becomes larger.
Gigas character occurs in both naturally as well as artificially-induced polyploids.
Physiological : Becker in 1931 reported that osmotic concentration in autopolyploids is
proportional top the chromosome number.
Autotetraploid maize takes longer time to mature than diploid. In fruits and vegetables,
vitamin content is increased. In tobacco, nicotine content is increased and also nitrogen,
calcium potas and magnesium but carbohydrate, sulphur and phosphorus decreases. In
triploid beet, sugar content is increased.

The decrease in growth rate slowly leads to the formation of perennial from annual types
in cotton. Sharp 1934 reported that autotetraploid Zea race is perennial, while diploid is
annual.
Polyploidy in Plants:Chromosome doubling in Somatic and Germ cells. Two basic irregular processes
have been discovered by which polyploids may evolve from diploid plants and become
established in nature.
1. With somatic doubling cells sometimes undergo irregularities at mitosis and give rise
to meristematic cells.
2. Reproductive cells may have an irregular reductional division in which the sets of
chromosomes fail to separate completely to the poles in anaphase. Both sets thus
become incorporated in the same restitution nucleus, which doubles the
chromosome number in the gamete.
3. Two main kinds of polyploids, autopolyploids and allopolyploids, may be
distinguished on the basis of their source of chromosomes. It occurs frequently in
single cells of many plants, but they not survive, Allopolyploids result when different
genomes come together through hybridization. Among the surviving plants.
The change in the chromosome:Number involve addition or elimination of individual chromosomes or of complete
chromosomal set (genome).on the basis of two main classes of ploids can be recognized1. Aneuploids or Heteroploids
2. Euploids
Numerical change in chromosomes or variation in the chromosome number
(heteroploidy) can be mainly of two types. Namely-Aneuploidy, Euploidy.

Aneuploidy means the presence of chromosome number, which is different than a

multiple of the basic chromosome number. Euploidy, on the otherhand, means that the
organism should possess one or more full sets of chromosomes, let us imagine that 7 is
the basic chromosome number (x) in a particular class of individuals where the diploid
number (2n), is 14. In this case, the chromosomes number 2n=15 and 2n=13 would be
aneuploids, while those having 2n =7,21,28,35 or 42 would be euploids. A classification
of different kinds of numerical changes in chromosomes is presented in fig-

10

Numerical changes in chromsomes

Euploidy

Aneuploidy

Monoploidy
diploidy Polyploidy
(x)
(2x)
(3x, 4x, 5x,6x etc)

hypoploidy

monosomy
(2n-1)

Trisomy

hyperploidy

nullisomy
(2n-2)

tetrasomy
(2n+1)

(2n+2)

Fig: Different kinds of numerical changes in chromosome (x=basic

chromosome number. 2n=somatic chromosome number).

11

Aneuploidy or Heteroploidy:It is a condition in which nuclei contain chromosomes whose number is not true
multiple of basic chromosome number or genome. The organism in which such
abnormal conditions exist are called Aneuploids.
Aneuploidy:The aneuploidy may be arranged in several groups, some of which are as follows:
1) Monosomic (2n-1):An organism lacking one chromosome of a diploid complement is called monosomic
(2n-1). Suppose in an organism diploid set is formed of 4 pairs of chromosomes. In this
by chance one chromosome is missing. So in this case one may fine 3 normal pairs of
chromosomes and one singlet chromosome. This should, there for, be called as
monosomic.
The monosomics are very rare. However, viable monosomics have been studied in
Drosophila, tobacco, Datura, Maize, tomatoes and many other organisms.
Monosomics show abnormal type of meiosis. They have chromosome constitution 2n-1,
so they produce 2 kinds of gametes; some with n and others with n-1 chromosome. The
odd chromosome, which has no homologue to pair tends to pass at random to one pole in
meiosis. Frequently, however, it acts as a laggard at anaphase and is not induced in any
of the daughter nuclei. For this reason the gametes having n chromosomes occur less
frequently than the gametes of (n-1) type. In plants, the nuclei with a missing
chromosome seldom survive. The n-1 gametophytic generation fails, presumably
because the complete absence of certain genes that occur on the missing chromosome
means the failure of certain biochemical reactions which are accomplished in presence

12

of enzyme formed by these genes. Thus monosomic analysis makes it clear what genes
are found on the particular chromosome or linkage group.
In some plants, e.g. Datura, monosomics are not viable. In Nicotiana tobaccum with
basic chromosome number 24, clausen and his co-workers have found all 24 possible
monosomics as expected and have noted that they were very much different from one
another. In wheat (triticum vulgare) haploid number n is 21. So one may expect that
21 monosomics of different types should be in bread wheat, but these do not differ
much from the normal diploids.
Nullisomic (2n-2):In these both the homologues of a particular chromosome set are somehow lost so
that a chromosomal pair is completely missing from the normal 2n set. The phenotypic
expression of such a plant, when compared with that of normal plant, will indicate the
number of genes contained in the missing chromosomes and the respective character
they governed. Nullisomics are generally nonviable like the monosomics. In maize and
wheat, several nullisomics have been reported. Nullisomics are generally obtained by
selfing monosomcis. When the gametes containing n-1 chromosomes fuse they form
nullisomics. Nullisomics are those individual, which lack a single pair of homologus
chromosomes, so that the chromosome formula would be 2n-2, and not 2n-1-1, which
would mean a double monosomic. Sears had isolated all the 21 nullisomics in wheat.
Trisomic (2n+1):An organism containing two complete genomes plus one extra chromosome is termed
trisomic (2n+1). In the normal process of meiosis, sometimes chromosomal pairs
separate in such a way that one member of each pair goes to one daughter nucleus and
other member goes to other daughter nucleus. But very rarely one pair of chromosomes
fails to divide and finally it moves as such to one spindle pole. So half of daughter cells
receive an extra chromosome and half of them lose one. In this way gametes with (n-1)
13

and (n+1) chromosomes will develop. If by chance the gamete with (n+1) chromosomes
fuses with normal gamete carrying n chromosomes, the resulting zygote will have
n+1+n or 2n+1 chromosomes and it will produce trisomic organism. Gametes with (n-1)
chromosomes on fusion with gamete carrying n chromosomes will give rise to
monosomic (2n-1) zygote. Many of the mutations in oenothera noted by Devries are
trisomics. Trisomis are widespread in nature and have been extensively studied in
Datura, maize, tomatoes, wheat, tobacco and Drosophilla.
Blakeslee and Belling (1924) announced a mutant type in Datura having 25
chromosomes rather than normal 24 chromsomes (2n=24). In that, at the meiotic
metaphase one of the 12 pairs was found to have an extra member, i.g. there were 11
normal pairs and one trisomic. The trisomic individulas exhibit several specific features
which are quite different from those seen in the wild types.
The additional traits were associated with the extra chromosome which gave to the
plants extra dose of all the genes that are already contained in the normal duplicated
chromosomes.
It is thus possible to identify some genes while are associated to extra chromosomes.
This supports the statement that genes are contained in particular chromosomes. Since in
Datura there are 12 chromosome pair with different genic composition and since the
triplication of every chromosome pair is possible,12distinguishable trisomics can be
expected in it. Blakeslee and his associates have succeeded in producing all 12 different
trisomics as expected. These trisomics were grown in Blakeslees garden and all were
found to have distinguishable phenotypic features which could easily be attributed to
extra dose of one chromosome to each of the 12 individual sets of chromosomes.
Trisomics show irregular meiosis. From a trisomic cell with total (2n+1)
chromosomes,the 2 kinds of gametes; some with (n+1) chromosomes, would be formed
after meiosis. Recessive genes present on the extra chromosomes in trisomic complexes
14

will express themselves less frequently than in normal diploid plants because
comparatively more wild type alleles are present to check their expression. It has been
observed by geneticists in Datura that extra chromosome enters the.
Production of Trisomics:Trisomics may originate spontaneously due to production of n+1 type of gametes
rarely as a result of non disjunction of a bivalent .

However:

2x

3x
Nondisjunction

n-1

n+1

2n+1

n+1

2n+1

Fig: Production of trisomics due to formation of n+1 type of gametes

in diploid (2x) and triploid (3x) individuals more aften trisomics are
produced artificially either by selfing the tripolids (produced by
crossing diploid and autotetraploids) or by crossing these triploids as
females with diploids as male (3x2x).
Cytology of trisomics:-

15

The trisomics have an extra chromosome which is homologous to one chromosome

of the complement. Therefore, it forms a trivalent this trivalent may take a variety of
shapes in primary and secondary trisomics as shown in fig. in a tertiary trisomic a
characteristic pentavalent is observed.
Type of trisomic
(a) Primary trisomics

Somatic chromosomes
1
1
1

Metaphase I configurations

111

2 2 2

1
(i)

(b)Secondary trisomics

(c) Tertiary trisomics

1
3
2
4

1 1

22

(i)

1 1

1 1

22
2 2 2

2 2

2 2

(ii)

2
2

1
(iii)

(ii)

11

2 2

(iv)

1 1

2 2

(iii)

3
2
2

1
1

4
4
(a pentavalent)

Fig: Different types of trisomics and their meiotic configurations at metaphase I .

Trisomic analysis:Trisomics are also used for locating genes on specific chromosomes. If a particular
gene is located on the chromosome involved in trisomy.
Double Trisomics (2n+1+1):-

16

3
3

Sometimes an organism receives two complete genomes plus two different extra
chromosomes. This is called double trisomic. The extra chromosomes are homologues of
two different chromosomal pairs in normal diploid set. Thus two different chromosome
pairs in the normal diploid set will have one additional homologue with them and thus
they form two trisomics. The phenotypic expression of genes on extra chromosomes in
trisomics will be deeper than in normal diploids.
Tetrasomic (2n+2):
Aneuploid nuclei which have normal diploid genome and one extra pair of
chromosomes are called tetrasomic (2n+2). In this one pair of chromosomes in the
diploid set is further reduplicated due to additional chromosome pair. Phenotypic
expression of the genes in tetrasome may be deeper than the affects found for the genes
in corresponding trisomics.
In meiosis the four homolouges of tetrasomic set after tend to form a quadrivalent. If
disjunction by twos is regular, a fair genetic system will operate. Generally quadrivalents
are not formed and disjunction

(Separation) is not always regular. Nevertheless,

tetrasomics are more regular than trisomics in meiosis.

Aneuploid segregation in plants:The first critical study of Aneuploid plants was initiated in 1924 by A.F. Blakeslee
and J. Belling when they discovered a mutant type with 25 chromosomes in the
common jimsonweed, Datura stramonium, which normally has 24 chromosomes in the
somatic cells. At the meiotic metaphase, one of the 12 pairs was found to have an extra
member, that is, one trisome (2n+1) was present along with 11 disomes. This trisomic
plant different from wild type plants in several specific ways, particularly in shape and
spine characteristics of seed capsules. Because the complement was composed of 12
chromosome pairs differing in the genes they carried, these were grown in Blakeslees

17

garden, each was found to have a distinguishable phenotype that was attributed to the
extra set of genes contained in one of the 12 chromosomes.
It is also possible to identify some traits that were determined by genes on particular
chromosomes by trisomic ratios that is, 5:1, 17:1 and 35:1, in contrast to 3:1 and 1:1
expected from monohybrid crosses in regular 2n plants. The extra chromosome in
poinsettia, for e.g. was found to carry the locus for alleles P + and P, for purple or white
flowers, respectively. Any one of these chromosome arrangements with the dominant
gene P+ (P+ P+ P+ , P+ P+P or P+ PP) produced poinsettia plants with purple flowers,
whereas only one, the fully recessive.
Uses of Aneuploids
1) They have been used to determine the phenotypic effects of loss or gain of different
chromosomes.
2) They are used to produce chromosomes substitution lines. Such lines yield
information on the effects of different chromosomes of a variety in the same genetic
background.
3) They are used to produce alien addition and alien substation lines. These are useful in
gene transfers from one species into another.
4) Aneuploid analysis permits the location of a gene as well as of the linkage group onto
a specific chromosome. This is one of the most important applications of aneuploidy.
5) Studies on nullisomic-tetrasomic combinations made it possible to establish
homoeology among the chromosomes of A, B and D genomes of wheat.
6) Multivalent formation in nullisomic 5B of wheat led to the discovery of the Ph gene.
This gene suppresses pairing between homoeologous chromosomes.
7) Aneuploids are useful in identifying the chromosomes involved in translocations.
8) Aneuploids are also useful in preparation of molecular maps.
9) They may be used for obtaining chromosome specific probes. A probe is a DNA
sequence that is used in nucleic acid hybridization for detecting the presence of the
same DNA sequence in test DNA samples.
Euploidy:18

(GK. Eu=true; and ploid=unit). This is a condition in which the change in the
chromosome number involves additions or elimination of complete genome.
The basic chromosome number of euploids are represented by the haploid (n). the
euploids above the haploid or monoploid level are called polyploids. They may be
diploid (2n), triploid (3n), tetraploid (4n), pentaploid (5n), hexaploid (6n) and so on. In
all these cases of euploids we see that the chromosome number are exact multiples of
their original haploid number (n).
Whereas aneuploids differ from standard 2n chromosome complements in single
chromosomes, euploids differ in multiples of n or x if n=x.
Monoploids (n) carry one genome. The n or x chromosome number is usual for gametes
of diploid animals,But unusual for somatic cell.Monoploidy is seldom observed in
animals, but is found in the male honeybee and other insects in which male haploids
occur.
By contrast, plants have gametophyte stage in their cycle that is characterized by the
reduced (n) chromosome number. In higher plants , this stage is brief and in
conspicuous, but in some lower plant groups it is the major part of the some lower plant
groups it is the major part of the cycle. Occasionally plants in natural populations or
plots can be recognized as monoploids by abservation and verified by cytological
producer. These plants are usually frail in structure with small leaves, low viability, and a
high degree of sterility. Sterility is attributed to irregularities at meiosis. Obviously, no
pairing is possible because only one set of chromosomes is present. Therefore, if the
meiotic process succeeds at all, the dispersal of chromosomes to the poles is irregular,
and the resulting gametes are highly in viable, because monoploids undergo no
segregation and carry a single set of gene.

19

It is several types. Some of them are as follows:1. Hapolyploidy or Monoploidy (a condition in which nucleus of an organism has only
single genome n).
2. Diploidy or 2n (in this condition, organisms have chromosomes in pairs, i.e. their
nuclei contain two complete genomes).
3. Polyplioidy (in this condition, organisms have more than two genomes per cell. For
example, triploids (5n), tetraploid (4n), pentaploid (5n), hexaploid (6n) and so on.
Haploids or Monoploids:Monoploids or haploid organisms contain single genome, i.e. they contain one
member of each kind of chromosome.
The haploid chromosome number (n) is usually found in gametes of diploid plants and
animals and this is unusual for a somatic cell of higher plant. Monoploidy has been
reported in several cases, both plants and animals. Majority of lower plants, particularly
thallophytes and bryophytes, exist in monoploid condition. In higher plants, haploids
develop as a result of parthenogenesis of haploid eggs. In Datura Stramonium, Oryza
sativa, Oenothera Triticum, Zeamays and many other plants, haploidy has been noted.
Characters of Haploids:
1) These plants are usually weak and small sized, but in pepper the haploids are as
healthy as normal diploid plants.
2) Leaves are generally small
3) Plants are of low viability
4) Cytology of haploids:High degree of sterility has been observed in these plants. It is so only because meiosis
is irregular. As the chromosomes in haploid set do not have their homologues, the
pairing of chromosomes is not possible and the chromosomes are found as univalent at
metaphase I of meiosis. So neither the eggs nor the pollens are formed. However, if
meiotic process succeeds at all, the univalent chromosome are found scattered all over
20

the cell. They may constitute restitution nucleus including all chromosomes and may
thus give rise to gametes having a complete haploid set of chromosomes. Sometime the
distribution of chromosomes to the two poles is irregular and the gametes so formed are
highly non-viable (i.e. they die soon).
Normal gamete is possible if by chance all the chromosomes pass to one pole during the
irregular meiosis. Monoploids do not undergo segregation and they carry a single set of
genes. So gemetically they are pure.
Use of Haploids:When the chromosome number of a haploid is doubled by colchicine treatment. The
derived diploid will be completely homozygous for all genes so that the most important
use of haploids is in the production of homozygous diploids. Seven varieties in rice,
three varieties in tobacco and two varieties in wheat have also been released using
haploids in china consult cytogenetic of crop plants.
Thus they can be used experimentally to a good advantage
Diploids:Most common type of euploid is the diploid which has two complete genomes. As
mentioned earlier this chromosome arrangement is nearly universal among animals and
common in plants. These organisms are normal in their behavior, so they are not
discussed here in detail.
Polyploids:Polyploidy is commonly met within plant world. In animals, it is indeed rare. About
one half of total species of flowering plants are polyploids (stabbins, 1950). Among
certain families the proportion of polyploids to diploids is higher than 50 percent. About
two-third species of all grasses are polyploids. The cases of polyploidy may be found in
many other common plants, e.g. Chryasanthemum, Solanum, Brassica, Wheat,
21

Nicotiana. In the genes chrysanthemum, the basic number of chromosomes is 9. In this

genes species are known with 36,54,63,72 and 90 chromosomes. In wheat the basic
chromosome number is 7. In this , several varieties are known with 14, 28 and 42
chromosomes which are diploid, tetraploid and hexaploid respectively. In the nicotiana
(12), varieties with 24, 48, 72 chromosomes are known.
Similarly the genus Solanam, in which basic chromosomes number is 12, consists of
several polyploids varieties with 24,36,48,60,72,96,108,120 and 144 chromosomes. In
Rose, the basic chromosome number, is 5. In this, diploid, triploid, tetraploid, hexaploid
and octaploids have 10,15,20,30 and 40 chromosomes respectively.
Types of Polyploids:Two features could account for the origin of polyploids from diploids.
1) Doubling of the chromosome number in somatic tissue capable of giving rise to
gametes.
2) Failure of reduction in the chromosome number during the formation of gametes. On
the basis of their origin, the polyploids are of the following two typesi) Autopolyploids:These individuals contain more than two identical genomic sets which are derived by
self-duplication of parental genomic sets.
In more scientific language, in these organisms, multiple genomes are identical or
very nearly so. Genome reduplication within a normal species give rise to
autopolyploids. It arises due to failure of anaphase during meiosis.
Because of these multiple sets, segregation problem may occur, such that the gametes
may not receive full sets of chromosomes and, as a consequence, the organism will
suffer reduced fertility.
For example, autotriploids (2n=3x have three complete chromosome sets. At any one
point only two chromosomes can pair effectively (although small sections of triple
22

paired synaptonemal complex have been seen in some triploids and trisomics. This
means that combinations can occur of three univalents, one univalent and a bivalent
or a trivalent. Univalents, will generally be lost from the cell via micronucleation.
Bivalents will segregate normally. The orientation of the trivalent on the metaphase
plate determines how many chromosomes go to each pole.
To ensure full fertility, all chromosomes must form trivalent, all trivalent must
orientate in a convergent manner and all convergent trivalents must segregate two
chromosomes to one pole and one to the other. This would produce two diploid
gametes and two haploid gametes. Needless to say, this rarely happens and triploids
form all types of configuration at metaphase I. this produces unbalanced gametes that
are generally non viable, and consequently fertility is reduced.
As a general rule, the higher the basis number, the lower the fertility. Other
polyploids with odd numbers of chromosomes sets 5x, 7x, etc. experience similar
problems to triploids because the odd set, which will either form univalents, trivalents
or complex multivalent. For example in a pentaploid when 2n=5x the chromosomes
can pair in any combinations of between one and five chromosomes.
When a triploid has a total chromosome number of 9, fertility is reduced to 25% of
the diploid level as a result of the probability of all chromosomes forming trivalents
and all trivalents orientating the same way. When 2n=3x=12, fertility is further
reduced to 12.5% compared with diploid. Higher basic numbers reduce fertility
further and are almost completely sterile. In triploid apples such as Blenheim Orange
and Bramleys Seedling, in which 2n=3x=51, seed set is less than 5% and the
viability of the seed set is well below 1OO%.
Even numbered polyploids such as tetraploids (2n=4x, in general, have a higher
level of fertility. Tetraploid chickpea has only a 15% reduction in fertility when
compared with the diploid.
23

Tetraploids have four complete sets of chromosomes. Each set of four homologues
has the following pairing options: four univalents; two bivalents; a bivalent and two
univalents; a univalents and a trivalent; or a quadrivalent. The frequency with which
these are produced depends on the pairing arrangements. The same basic rule as
described for triploids also applies here: segregation of complete sets of
chromosomes produces viable gametes; other combinations either produce non viable
gametes, or result in viable or deleteriously affected offspring.
Many tetraploids have mostly bivalent and quadrivalent pairing as this allows for
maximal pairing in the synaptonemal complex, producing balanced segregation at
anaphase I.
Cytology of autopolyploids:In an autopolyploids, there will be more than two sets of homologous chromosomes.
This leads to the formation of multivalent instead of bivalents as found in the diploids. In
an autotriploids there three sets of homologous chromosomes.If these three sets are
normally paired, trivalent as described in primary trisomics will be observed. The
trivalents can not disjoin normally and will either disjoin 2:1 chromosomes to two pales
or will disjoin 1:1 leaving one chromosomes as a laggard. The number of chromosomes
in the gametes of a triploid organism therefore, will vary from n to 2n. Most of these
gametes are unbalanced leading to high degree of sterility.
In autotetraploids, since there are four sets of chromosomes quadrivaents are formed
which disjoin in a normal 2:2 manner giving diploid gametes. Rarely, however, a
quadrivalent may disjoin 3:1 or may leave one or more chromosomes, as laggards at
anaphase I. Therefore, autotetraploids also have certain degree of sterility, although it
will not be as high as in autotriploids.
Genetics of autopolyploids (trisomic and tetrasmoic inheritance):The segregation pattern in polyploids is quite different than what we find in diploids.
24

In a polyploidy there are more than one kind of heterozygotes, because there are more
one kind of heterozygotes, because there are more than two homologues. The different
possible genotypes and expected the ratios are based on the assumption that the gene is
close to centromere leading to what we cell chromosome segregation. If gene is away
from centromere, crossing occur between gene and the centromere will take place and
the ratios will be modified. This will then be called chromatid segregation, because
segregation is taking place not at the chromosome level but at the chromatid level due to
crossing over.
Autotriploids:Such plants have three genomes per nucleus and they are commonly designated as 3n
plants. Autotriplids occur in a number of plants, e.g. Oenothera, Datura, rose, rice and
many others. They are usually formed as a result of fertilization between a diploid (2n)
and a haploid gamete (n), (2n gamete+ n gamete=3n zygote).
Diploid gametes are produced by normal tetraploids in meiosis or in sectors of otherwise
diploid organisms where automatic doubling of somatic chromosome number has taken
place. Somatic doubling may be spontaneous or may be induced.
Characters:i)Autotriploids are more vigorous than normal diploids, they are more leafy and show
ii)
iii)

tendency towards perenniality.

Sometimes floral abnormalities may be observed.
Pollengrains, stomatal guard cells and wood cells of xylem in triploids are larger

iv)

than those of diploids.

Plants are highly sterile and seeds are formed rarely. In nature, seed propogated
triploid plants are ordinarilly by vegetative means this is advantageous in
horticulture, specially for ornamental plants, e.g. chrysanthenum roses. Dahlias etc.
and also in the production of seedless fruits, some of the commercially important

25

fruit plants, e.g. apples, pears, bananas, grapes, orange, guavas, pineapples are
triploids.
Cytological behavior of Autotriploids :In autotriploid it is irregular. The centromeres of three homologous chromosomes
have no way to orient themselves so as to give equivalents at the two poles. In this
condition, the only possibility is that the components of trivalent separate in such a way
that two chromosomes go to one side and third goes to other. If it is so then the meiotic
products may contain either n or 2n chromosomes or any number in between. This is
true irrespective of whether three homologous chromosomes align either as trivalent or
as one bivalent and one univalent. Therefore, the gametes arising from triploids, have
unbalanced genomes. In fact, probability of formation of haploid and diploid gametes
are vary rare.
Autotetraploids-4n:Autotetraploids have four similar genomes per nucleus. They may originate in one of
the following ways.
i)By fusion of two diploid gametes.
ii)
They may result from the duplication of somatic chromosomes (2n) and following
failure of mitotic division. The resulting nucleus contains 4 copies of each
chromosomes, instead of the usual two. It this tetraploid perpetuates itself through
normal mitosis, the increased chromosome number may become established in a
group of cells or tissues within the body of organism. Plants capable of vegetative
propagation may be manipulated to produce pure tetraploids.
Characters:i)Autotetraploids usually show marked phenotypic variations, sometimes marked disease
resistance, they are commercially more valuable than corresponding diploids.

26

ii)

Some are larger and more deeply coloured than diploids and sometimes they bear

iii)

large seeds and fruits.

They have larger cells and epidermal cells are larger and are arranged in different

iv)

patterns in tetraploids than those in diploids.

The stomatal guard cells and epidermal cells are larger and are arranged in different

v)
vi)
vii)

patterns in tetraploids than those in diploids.

Flowers are larger and more showy in tetraploids than those in diploids.
High content of vitamin C (ascorbic acid) have been reported in tetraploids.
The tetraploids show tendency towards perenniality and may show low fertility.

Cytological behavior in Autotetraploids:In autotetraploids, each chromosome is present four times. Thus, chromosome
associations such as quadrivalents, trivalents, bivalents and univalients are expected
based on random association of four homologus chromosomes. Chromosomes pairing is
usually studied at diakinesis or metaphase I of meiosis, because chromosomes at these
stages are in condensed forms, chromosome migration at anaphase-I in autotetraploids is
determined by the co-orientation of their kinetochores at metaphase I.
It may be normal or abnormal. When four homologues align in tetravalents, normal
diploid gametes are formed. Sometimes, two bivalent or one trivalent and one univalent
may also occur. In these cases, products of meiosis are irregularly formed. Incomplete
genomes in gametes may result in total sterility.
Allopolyploids:Alloployploids are obtained when crosses are made between two member of distinct
taxonomic groups. The hybrid so obtained are highly sterile because the genomes
coming from two different individuals are so different that their chromosomes do not
synapse during zygote stage of meiosis I. These hybrid allopolyploids owing to complete
sterility may face extinction unless they are able to reproduce vegetatively. As far as
meiosis is concerned, the situation in allopolyploids is exactly like that in haploids.
27

Regular meiosis and formation of gametes containing one complete genome from each
of the two parents are possible only if the number of chromosomes in somatic cells is
doubled.

When such gametes fuse they give rise to a zygote containing the complete diploid
complement of original parents. Double diploids of this sort are called amphidiploids.
They are fertile, since each chromosome has now got its pairing partner. These
amphidiploids behave normally like a diploid with relatively large number of
chromosomes. The origin of amphidiploids is explained in the figure.
A very important case of allopolyploid is Raphanobrassica which was experimentally
evolved by the Russian geneticist G.D. Karpechenko who made intergeneric cross
between radish (Raphanus sativus) and cabbage (Brassica oleracea). Although these
plants were distantly related they were enough alike to be crossed successfully with each
other. Both had 9 pairs of chromosomes. The diploids hybrid Raphanobrassica had 18
28

chromosomes. 9 chromosomes from radish and 9 chromosomes from Brassica parent.

But it could not perpetuate itself largely because of failure of pairing between unlike
chromosomes during meiosis. When chromosomes of sterile Raphanobrassica were
doubled by artificial means a fertile polyploid Raphanabrassica was produced with 36
chromosomes; 18 of radish and 18 of cabbage. Thus the homologous chromosomes
could pair themselves and the regular meiosis resulted normal gametes, each with 18
chromosomes. This pairing of chromosome is called autosynapsis, as actually
chromosomes of two genomes are pairing with their respective replicas. This experiment
had theoretical significance since it provided a method by which fertile inter specific
hybrids could be produced. This also suggests the possibility of incorporating the
desirable genotypes of two different species into a new polyploid species.
Radish (Raphanus Sativus) Cabbage (Brassica Oleracea)
(2n=18)

(2n=18)
F1 hybrid sterile
(n = q + n = q)
The allopolyploidy thus represents the method by which new species originate in nature.
Fruits of amphidiploids are characteristically someuehat larger and plants as whole
possess a good deal of the robustness.

29

30

Evolution of new species due to allopolyploidy is seen among the cultivated verities of
wheat. Cytogenetically analysis of cultivated wheat plants has yielded important
information in this regard. Wheat verities fall under three groups: diploid, allotetraploid
and allohexapeoid. These are given in the following table.
The genome C present in bread wheat is supposed to have been derived form a grass
aegilops squrarrosa (2n=14) which grows in the region extending from Armenia to
Afghanistan. The different varities of wheat with higher chromosome number have
apparently arisen from this type and other related grasses through hybridlization
followed by chromosome doubling.
The individuals in which the genomes making a multiple set are not alike are called
allopolyploids. The different genomes are derived from two or more distinct species by
hybridization. Thus, it involves addition of new genomic sets from outside.
Many even- numbered polyploids may be allopolyploids with little similarity between
the different contributing chromosome sets. Thus, in order to maintain fertility, bivalents
are only formed between homologous chromosomes, for example the hybrid formed
between Brassica oleracea (2n=2x=18 and Raphanus sativus (2n=2x=18. The F1 is
sterile owing to lack of pairing between the two chromosomes sets, but if these are
doubled the allotetraploid (2n=4x=36 is fertile with the formation of 18 bivalents.
Bivalents may also be formed in even- numbered polyploids owing to the effects of
various pairing control mechanism (such as Pb in wheat which suppress homoeologous
pairing.
The addition of B chromosomes to polyploid cell can also increase the rate of bivalent
formation of approximately 15% when compared with the same variety without Bs.
Some specific Examples of Autopolyploids:

31

A) Evolution of wheat:Common cultivated wheat is another example of allopolyploidy, although its

allopolyploid nature has now been questioned. There are three different chromosome
numbers in the genus Triticum, namely 2n = 14, 2n = 28 and 2n = 42. The common
wheat is hexaploid with 2n = 42and is derived from three diploid species: AA = Triticum
aegilopoides (2n = 14), (ii) BB =Aegilops speltoides, (2n = 14) and (iii) DD = Aegilops
squarrosa (2n = 14).The hexaploid wheat, therefore, is designated as AABBDD, the
tetraploid (2n = 28) as AABB. and diploid (2n = 14) as AA. There is, however, evidence
available now which suggests that A, B &D genomes from three diploid species.
B) Synthesized allopolyploids:Certain allopolyploids were artificially produced in order to find out the origin of
naturally occurring allopolyploids. Common hexploid wheat and tetraploid cotton
furnish two such examples.
i)Tritium spelta, a hexaploid was artificially synthesized in 1946 by E.S. McFadden and
E.R. Sears and also by H. Kihara. They crossed an emmer wheat (tetraploid; 2n=28)
with Aegilops squarrosa (diploid; 2n=14) and double the chromosome number in
the F1 hybrid. The hexaploid or amphidiploid synthesized in this manner was found
to be similar to the primitive wheat T. spelta when this synthesized hexaploid was
crossed with naturally occurring T. spelta, the F1 hybrid was completely fertile and
showed normal pairing of chromosomes into bivalents. This suggested that
hexaploid wheat must have originated in the past due to natural hybridization
between tetraploid wheat and goat grass (Aegilops squarrosa) followed by
subsequent chromosome doubling.

Triticum dicoccoides
(tetraploid wheat)

32

Aegilops squarrosa
(goatgrass)

AABB
2n=28; 14 bivalents

DD
2n=14; 7 bivalents

ABD
2n=21; 21 univalents (triploid hybrid)
colchicine

AABBDD
2n=42; 21 bivalents
synthesized hexaploid wheat
(Triticum spelta)
Fig: Artificial synthesis of hexaploid wheat.
ii) Gossypium hirsutum, popularly known as upland cotton, is another interesting example
of amphidiploidy. Old world cotton has 13 pairs of larg chromosomes (genome Ah),
while American or upland cotton has 13 pairs of smaller chromosomes (genome
Dh). The new world cotton, the cultivated long-staple type has 26 pairs of
chromosomes (Ah Ah Dh Dh), 13 large and 13 small, J.O. Beasley crossed the
American and old world cottons and double the chromosome number in the F1
hybrids. The amphidiploid thus produced resembled the cultivated new world cotton
and when crossed with it gave fertile F1 hybrids. Thus, showed that New world
cotton, cultivated gossypium hirsutum (tetraploid) originated from two diploid
species namely G. herbaceum (2n=26) and G. raimondii (2n=26).

33

Gossypium herbaceum

Gossypium raimondii
Old world
American (upland)
Cotton
cotton
2n=26; 13 bivalents
2n=26; 13 bivalents
(Small chromosomes)
(large chromosomes)
F1 hybrid
2n=26; 26 univalents
(13 Small + 13 large)
Colchicine
2n=52; 26 bivalents
(13 small + 13 large)
New world cotton
(Gossypium hirsutum)
Fig: Artificial synthesis of New world cotton.
iii)Spartina townsendii, commonly called as Townsends grass and found in Europe, has a
chromosome number, 2n=126. This species in its morphology is intermediate to S.
altermi flora (2n=70), the American marsh grass and Spartina stricta (2n=56), the
European marsh grass. The study of the morphology and cytology of these three
species and the fact that S. townsendii was fertile suggested that Spartina townsendii
originated by the natural hybridization of S. altermifora and S. stricta, followed by
doubling of chromosome number in the F1 hybrid so derived. This is thus an
amphidiploids.

Spartina alterniflora
Spertina stricta
2n=70; 35 bivalents
2n=56; 28 bivalents

34

F1 hybrid
2n=63; 63 univalents
chromosome
doubling
2n=126; 63 bivalents
(S. townsendii)
Fig: Probable origin of Spartina townsendii.

35

C) Triticale, a new man made cereal:In recent years, considerable emphasis has been laid on the possibility of utilizing a
new man made cereal known as triticale (X Triticose cale wittmack). on a commercial
scale. It is already grown on about one million hectares (25 lakh acres) of land
commercially and research work is in progress all over the world to improve this man
made crop.
Triticale is the first man made crop, in so far as it resulted as an artificial allopolyploid
derived by crossing wheat (tritium) and rye (secale). Depending upon whether, tetraploid
(2n=4x=28) or hexaploid tritium (2n=6x=42) is utilized for the synthesis, one would get
hexaploid triticale (2n=6x=42) or octoploid triticale (2n=8x=56) respectively.
In each case, only diploid rye (2n=2x=14) was used. The derivations of triticales are
shown in fig.
In a similar manner, tetraploid triticales (2n=28) at some places were obtained, by using
diploid wheats. Decaploid triticale, using hexaploid wheat (2n=6x=42) and tetraploid rye
(2n=4x=28) were also obtained by A.Muntzing in Sweden. However, at present only the
hexaploid triticales (2n=42) are considered to hare the potentiality of becoming a new
crop. These hexaploid triticale are being crossed to wheat for making improvement.
As a result of these crosses a large no. of Secondary triticales have now been derived
from the original Primary triticales some of these secondary triticales have been,
released as commercial cultivars. Some of the most significant work on triticales was
done by A. Muntzing of Sweden, who passed away on 7 th January 1984. For a more
detailed account on triticales, readers are advised to consult either the book TriticaleResults and Problems written by A Muntzing in 1979.

36

(a)

Triticum durum
2n=28

Secale cereal
2n=14

F1 hybrid (Sterile)
2n=21
Chromosome doubling
2n=42
Hexaploid triticale
(b)

Triticum aestivm

Secale cereal
2n=42
2n=14
F1 hybrid (sterile)
2n=28
Chromosome doubling
2n=56
Octoploid tritcale

Fig: Artificial synthesis of (a) hexaploid triticale and (b) octaploid

triticale.

37

Varieties

Diploid chromosomes

Genome constitution

number
1. Small

grained

2n=14

AA

2n=28

AA BB

Einkorn types or
wheat of Europe
and Asia.
2. Emmer wheat of
Northern
which

Europe

has

Allotetraploid

thick

kernels and is used

mainly

as

stock

feed.
3. Bread

wheat

or

2n=42

vulgar wheat

AA BB CC
Allohexaploid

By colchicine treatment in order to get diploid gametes from these two plant species. By
crossing these two colchicine treated plants they found a fertile hybrid with 42
chromosomes. The hybrid resembled bread wheat and was given name T.spelta. These
researches showed how a moderately useful wheat and useless grass were combined in
nature to produce most valuable variety of wheat.
Application of Polyploidy:Among the cultivated varities of wheat, three different chromosome number are
represented: 14,28 & 42 (x=7) for example:

38

The primitive small grained einkorn type of Europe and Asia. Triticum
monococcum, has 14 chromosomes in its vegetative cells. Its yield is low and it is of
comparatively little value.
The bread wheats, T. aestivum, with 42 chromosomes, were postulated by
J.Percival in England to have come from a cross between emmer wheat and goat grass,
both of which are native to the Babylonian region where bread wheat originated.
Wheat techniques for artificial chromosome doubling became established,
investigations of the origin of bread wheat confirmed Percivals theory. Experimental
evidence obtained by E.S. McFadden and E.R. Sears and Separately by H. Kihara traced
the path way for the origin of one type of bread wheat.
The tetraploids (n=14), T. dicoccum and T. dicoccoides. Hybrids between these
tetraploid species of wheat and the hybrids were completely or nearly sterile when the
F1 hybrids of T. dicoccoides XA. squarrosa were treated with colchicine, highly fertile
alloploids with 42 chromosomes were obtained. These synthetic hexaploids closely
resembled the cultivated t. spelta and they produced highly fertile hybrids with that
species and with t. vulagre, known to be in the genome of the hexaploid wheats
contained one chromosome set that originated from A. squarrosa. It was postulated that
T. spelta is the ancestral hexaploid wheat of Europe, having arisen, possibly in fairly
recent times, in south eastern Europe.
New World Cotton:Crosses can be made between distinct species of cotton members of the genes
Gossipium. The hybrids show a wide range of rigor and fertility, making the material
favorable for studies of origins.
Old world cotton has 13 pairs of large chromosomes. American cotton has 13
pairs of small chromosomes. New world cotton has 26 pairs, 13 large and 13 small.

39

Evidently, hybridization and chromosome duplication occurred somewhere in the

ancestry of the new world cotton.
Primrose Hybridization:The primo some primula kewensis, is an allotetraploid with 36 (2n) chromosomes. It
was derived from a cross between two diploids. P. floribunda (x=9) and P. verticillata
(x=9). Plants from these two species crossed readily. Producing hybrids with 18
chromosomes in their vegetative cells, but the hybrids were sterile, the fertile
allotetraploid are shown.
Desease Resistance in Tabacco:Induced polyploidy has been exploited to a great extent. Practical applications may
become more common as additional data are accumulated.
By artificially induced polyploidy, disease resistance.
Polyploid Fruits, Flowers and wheat:Some varities of plants that serve human needs more effectively than other have now
been identified as polyploids.
Many polyploids were selected and cultivated because of their large size, vigor,
and ornamental values, before their chromosome numbers were know. Giant sports
from twigs of Mclntosb apple trees that were found to be tetraploid (4n) were propagated
into whole trees, which produce extra large fruit. Bartlitt pears, several varieties of
grapes, and cranberries have also produced sports with giant fruits. Some of these show
promise of practical usefulness, with colchicine treatment, a number of polyploids have
been developed artificially. This technique has provided a way to explore the mechanism
involved in polyploidy formation, and to make use of the good qualities of opolyploids.
Tetraploid (4n) maize is more vigorous than the ordinary diploid and produces some 20
percent more vitamin A. its fertility is somewhat reduced. Polyploid watermelons have
40

been developed from colchicine treatment by kihara and others. The tetraploid with 44
chromosomes is large and has practical value.
A constant threat to the wheat crop is rust a fungus that attacks the stems and leves
of the growing plants and destroys the ripening grain. Spores are borne by wind and,
when conditions are right, the disease can be combated by developing rust-resistance
strains.
In plant breeding:Various cereal crops and vegetables with several desirable characters have been
evolved by inducing polyploidy and crossing the polyploids. The polyploidy plants are
better in quality, taste, yield, nutritional value and sugar contents. Many polyploids are
disease resistance. Thus the knowledge of polyploidy is of great help in plant breeding.
Many species of wheat are polyploids. There are three categories in wheat, namely
Einkorm group (2n=14). Vulgare wheat (2n=42). The third verities is polyploidy and is
better than any of the other two groups, in Solanum and Nicotiana many good varities
could be evolved artificially by the knowledge of polyploidly. Hybrids which are proved
useless in the beginning, the application of polyploidy inducing agents made it possible
to double the chromosome number in the sterile hybrids.
The genome analysis shows that first species is amphidiploid hybrid of the last two
species. Nicotiana tabaccum is susceptible to downy mildew disease but N. glauca, an
Australian species, is disease resistance polyploidy which contains more nicotine content
and shows more vigour.
Among ornamental varieties, certain triploids are of outstanding attraction in comparison
to diploids because of their large flowers, longer flowering period. Some of the best
varieties of apples are triploids.

41

Role of Polyploidy in Plant:The polyploidy has played an important role in evolution of new varieties in nature.
Angiosperms and pteridophytes have very high number of polyploid species in nature.
More than 35% angiosperms are polyploids. According to Manton (1950) among all the
plant groups, ferns show highest degree of polyploidy. It is generally noted that with the
increase in chromosome number the adaptability and variabilities of species increase
progressively. Many mutation in diploids are deleterious or fatal, but in polyploids,
where a gene is represented more than twice the chances of appearance of deleterious
effects due to mutation are lesser.

42

CONCLUSION
Polyploidy is the heritable condition of possessing more than two complete sets of
chromosomes. Most polyploids have an even number of sets of chromosomes, with four
being the most common (tetraploidy). Polyploids are very common among plants and
common among fish and amphibians, and are usually fit and well adapted. Indeed, the
study of eukaryotic genomes is providing surprising proof of the evolutionary potential
of polyploids: many sequenced genomes display the signature of polyploidy ancestry1
8. This indicates that polyploidy can bestow long-term evolutionary flexibility, instead of
freezing species in a static state that is enforced by gene redundancy, as was originally
proposed the phenotypic and molecular characterization of NEOPOLYPLOIDS, it has
been inferred that after polyploids form they pass through a bottleneck of instability
before becoming adapted and joining the evolutionary fray as efficient competitors of
their diploid relatives. Adapted polyploids that avoid extinction enter an evolutionary
trajectory of DIPLOIDIZATION, during which genomic redundancy is reduced14,15.
Duplicated genes can be lost, retained or maintained as duplicates, often undergoing
SUBFUNCTIONALIZATION and NEOFUNCTIONALIZA TION16,17. Bioinformatic
and theoretical analyses indicate that these processes are often not random and that the
function and properties of the encoded protein affect the outcome1825. By providing
duplicated genes, polyploidization might fuel long-term diversification and evolutionary
success.
In this review I discuss possible advantages of polyploidy and constraints on polyploid
formation that are indicated by either experimental evidence or theoretical
considerations. Becoming and remaining polyploid changes the organization and
function of the genome at both genetic and EPIGENETIC levels. For example, in
addition to the creation of gene redundancy,

43

polyploidy causes nuclear enlargement and increases the complexity of the processes
that are involved in managing and partitioning chromosomes during cell division.
Perhaps the most striking evidence of change comes from the discovery of epigenetic
remodelling, which leads to both the activation and suppression of gene expression.
Although some of these changes are potentially advantageous, many cause instability of
the neopolyploids and might be disruptive. This review of how polyploids form and
adapt will not only be useful to readers who are interested in the evolutionary role of
polyploids. It will also be useful to those who are interested in how parental gene
interactions can lead to HETEROSIS or DYSGENESIS, in how epigenetic patterns are
maintained or altered, and in how structural features of genomic organization affect
cellular functions.

44