“Introduction to Molecular

Biology”
Assignment # 01

DEPARTMENT OF BIOSCIENCES

CIIT ISLAMABAD

Polyploidy
Polyploidy refers to a numerical change in the whole set of chromosomes.
Polyploid cells and organisms are those containing more than two paired
(homologous) sets of chromosomes. Most species whose cells have nuclei
(Eukaryotes) are diploid, meaning they have two sets of chromosomes—
one set inherited from each parent.

In other words, the polyploid cell or organism has three or more times the

haploid chromosome number. Polyploidy arises as the result of total
nondisjunction of chromosomes during mitosis or meiosis. Polyploidy is
common among plants and has been, in fact, a major source of speciation
in the angiosperms. For example, triploid (3n) and tetraploid cell (4n) cells
are polyploid.
In addition, polyploidy occurs in some tissues of animals that are otherwise
diploid, such as human muscle tissues.  This is known as endopolyploidy.
Species whose cells do not have nuclei, that is,Prokaryotes, may be
polyploid organisms, as seen in the large bacterium Epulopiscium
fishelsoni . Hence ploidy is defined with respect to a cell. Most eukaryotes
have diploid somatic cells, but produce haploid gametes (eggs and sperm)
by meiosis. A monoploid has only one set of chromosomes, and the term is
usually only applied to cells or organisms that are normally diploid.
Male bees and other Hymenoptera, for example, are monoploid. Unlike
animals, plants and multicellular algae have life cycles with two alternating
multicellular generations. The gametophyte generation is haploid, and
produces gametes by mitosis, the sporophyte generation is diploid and
produces spores by meiosis.

Polyploidy may occur due to abnormal cell division, either during mitosis, or

commonly during metaphase I in meiosis.
Polyploidy occurs in highly differentiated human tissues in the liver, heart
muscle and bone marrow. It occurs in the somatic cells of someanimals,
such as goldfish,  salmon, and salamanders, but is especially common
among ferns and flowering plants (see Hibiscus rosa-sinensis), including
both wild and cultivated species. Wheat, for example, after millennia
of hybridization and modification by humans, has strains that
are diploid (two sets of chromosomes), tetraploid (four sets of
chromosomes) with the common name of durum or macaroni wheat,
and hexaploid (six sets of chromosomes) with the common name of bread
wheat. Many agriculturally important plants of the genus Brassica are also
tetraploids.
Polyploidy can be induced in plants and cell cultures by some chemicals:
the best known is colchicine, which can result in chromosome doubling,
though its use may have other less obvious consequences as
well. Oryzalin will also double the existing chromosome content.

Types of Polyploidy:

There are two general types of ploidy, which include plants that have either
one or more complete sets of chromosomes present in their genome
(euploids) or those that have partial sets due to the absence of at least one
of their individual chromosomes or presence of at least one extra one
(aneuploids):

1. Euploidy:
Individual has one or more whole sets of chromosomes with each
chromosome represented once in each set. Following are 3 main
kinds of euploidy.
 Haploidy: Individual with half (n=x) of the somatic cell chromosome
number.
 Diploidy:Iindividual with two sets of the basic, complete genome
(2n=2x).
 Polyploidy: Individual with more than two basic, complete sets of
chromosomes in its somatic cells. Among polyploids, these are main
types:
  Autoploidy: Individual has more than two complete chromosome
sets from a single genome. Also known as autopolyploidy.
 Alloploidy:Iindividual has two or more genomes contributed
from different parental species in their ancestral lineage. Also known
as allopolyploidy. Most naturally occurring polyploids are alloploids.
  Amphidiploids: (Also called amphiploids) are allotetraploids that
contain two sets of genomes from each of two different parents.
Amphidiploids are so called because they behave like diploids during
meiosis.

2. Aneuploidy:
Organism has a partial set of chromosomes due to addition or
deletion of specific chromosome(s). or one or more chromosome
sets. It is a state in which the number of chromosomes is not an exact
multiple of the haploid number.

Origin of Polyploidy:
Polyploidy has occurred often in the evolution of plants.

The process can begin if diploid (2n) gametes are formed. These can
arise in at least two ways.

 The gametes may be formed by mitosis instead of meiosis.

 Plants, in contrast to animals, form germ cells (sperm and eggs) from
somatic tissues. If the chromosome content of a precursor somatic
cell has accidentally doubled (e.g., as a result of passing through S
phase of the cell cycle without following up with mitosis and
cytokinesis), then gametes containing 2n chromosomes are formed.

Polyploidy also occurs naturally in certain plant tissues.

 As the endosperm (3n) develops in corn (maize) kernels (Zea mays),

its cells undergo successive rounds (as many as 5)
of endoreplication producing nuclei that range as high as 96n.
  When rhizobia infect the roots of their legume host, they induce the
infected cells to undergo endoreplication producing cells that can
become 128n (from 6 rounds of endoreplication).

Polyploidy can also be induced in the plant-breeding laboratory by treating

dividing cells with colchicine. This drug disrupts microtubules and thus
prevents the formation of a spindle. Consequently, the duplicated
chromosomes fail to separate in mitosis. Onion cells exposed to colchicine
for several days may have over 1000 chromosomes inside.

Polyploidy and Speciation:

When a newly-arisen tetraploid (4n) plant tries to breed with its ancestral
species (a backcross), triploid offspring are formed. These are sterile
because they cannot form gametes with a balanced assortment of
chromosomes.

However, the tetraploid plants can breed with each other. So in one
generation, a new species has been formed.

Polyploidy even allows the formation of new species derived from different
ancestors.

In 1928, the Russian plant geneticist Karpechenko produced a new species

by crossing a cabbage with a radish. Although belonging to different genera
(Brassica and Raphanus respectively), both parents have a diploid
number of 18. Fusion of their respective gametes (n=9) produced mostly
infertile hybrids.

However, a few fertile plants were formed, probably by the spontaneous

doubling of the chromosome number in somatic cells that went on to form
gametes (by meiosis). Thus these contained 18 chromosomes — a
complete set of both cabbage (n=9) and radish (n=9) chromosomes.

Fusion of these gametes produced vigorous, fully-fertile, polyploid plants

with 36 chromosomes. (They had the roots of the cabbage and the leaves
of the radish.)
These plants could breed with each other but not with either the cabbage or
radish ancestors, so Karpechenko had produced a new species.

The process also occurs in nature. Three species in the mustard family
(Brassicaceae) appear to have arisen by hybridization and polyploidy from
three other ancestral species:

 B. oleracea (cabbage, broccoli, etc.) hybridized with B. nigra (black

mustard) → B. carinata (Abyssinian mustard).
 B. oleracea x B. rapa (turnips) → B. napus (rutabaga)
 B. nigra x B. rapa → B. juncea (leaf mustard)

Modern wheat and perhaps some of the other plants listed in the table
above have probably evolved in a similar way.
Polyploidy in Animals:
Although polyploidy can be of benefit for organisms, there are many fewer
species of polyploid animals than plants. The exact reason for this is not
entirely known. Some scientists think that it might have to do with the
increased complexity of animal body plans compared to plants. Other
suggests polyploidy may interfere with gamete formation, cell division, or
regulation of the genome. However, there are some exceptions in fish,
reptiles, and insects.

The New Mexico whiptail is an example of a triploid animal

New Mexico Whiptail:
A specific example of polyploidy in action is the triploid trout. In Idaho, the
cutthroat trout is the state fish, but other fish, like the rainbow trout were
introduced to the area.

conservationists wanted to prevent the rainbow trout from breeding with the
cutthroat trout, to preserve their species. The solution? Create a triploid
rainbow trout that can't reproduce using polyploidy.

So how did scientists manage this? It turns out that fish reproduce slightly
differently than humans. Eggs start as diploid, and sperm as haploid. When
the cells fuse, the new cell kicks out the third set of chromosomes, so the
new fish can be diploid. However, if the last set of chromosomes isn't
kicked out, the new fish will become triploid, and unable to reproduce.
Scientists took advantage of this by exposing the fish eggs to warm water
in a hatchery, preventing the extra chromosomes from being kicked out.
The now triploid and sterile fish could then be introduced to the river
without fear of them mating with the local cutthroat population.

Examples in animals are more common in non-vertebrates such as

flatworms, leeches, and brine shrimp. Within vertebrates, examples of
stable polyploidy include the salmonids and many cyprinids (i.e. carp Some
fish have as many as 400 chromosomes.Polyploidy also occurs commonly
in amphibians; for example the biomedically-important Xenopus genus

contains many different species with as many as 12 sets of chromosomes

(dodecaploid.Polyploid lizards are also quite common, but are sterile and
must reproduce by parthenogenesis.[citation needed] Polyploid mole
salamanders (mostly triploids) are all female and reproduce by
kleptogenesis. "stealing" spermatophores from diploid males of related
species to trigger egg development but not incorporating the males' DNA
into the offspring. While mammalian liver cells are polyploid, rare instances
of polyploid mammals are known, but most often result in prenatal death.

Polypolidy in Humans:
Polyploidy occurs in humans in the form of triploidy (69,XXX) and
tetraploidy (92,XXXX).
Triploidy occurs in about two to three percent of all human pregnancies
and around 15 percent of miscarriages. The vast majority of triploid
conceptions end as miscarriage and those that do survive to term typically
die shortly after birth. In some cases, survival past birth may occur longer if
there is mixoploidy, with both a diploid and a triploid cell.
Tripolgy may be the result of either diandry (the extra haploid set is from
the father) or digyny (the extra haploid set is from the mother). Diandry is
almost always caused by the fertilization of an egg by two sperm
(dispermy). Digyny is most commonly caused by either failure of one
meiotic division during oogenesis leading to a diploid oocyte or failure to
extrude one polar body from the oocyte.
Diandry appears to predominate among early miscarriages, while digyny
predominates among triploidy that survives into the fetal period. However,
among early miscarriages, digyny is also more common in those cases
under 8.5 weeks gestational age or those in which an embryo is present.
There are also two distinct phenotypes in triploid placentas and fetuses that
are dependent on the origin of the extra haploid set. In digyny, there is
typically an asymmetric poorly grown fetus, with marked adrenal hypoplasia
(incomplete or arrested development of the adrenal glands) and a very
small placenta. In diandry, the fetus (when present) is typically normally
grown or symmetrically growth restricted, with normal adrenal glands and
an abnormally large cystic placenta that is called a partial hydatidiform

mole. These parent-of-origin effects reflect the effects of genomic

imprinting.
Complete tetraploidy is more rarely diagnosed than triploidy, but is
observed in one to two percent of early miscarriages. However, some
tetraploid cells are not uncommonly found in chromosome analysis at
prenatal diagnosis and these are generally considered "harmless." It is not
clear whether these tetraploid cells simply tend to arise during in vitro cell
culture or whether they are also present in placental cells in vivo. There
are, at any rate, very few clinical reports of fetuses/infants diagnosed with
tetraploidy mosaicism.
Mixoploidy is quite commonly observed in human preimplantation
embryos and includes haploid/diploid as well as diploid/tetraploid mixed cell
populations. It is unknown whether these embryos fail to implant and are
therefore rarely detected in ongoing pregnancies or if there is simply a
selective process favoring the diploid cells.
Polypolidy in Plants:
Scientists have hypothesized that two thirds of flowering plants are
polyploid. Most ferns and grasses are polyploid, and even crops that we
enjoy everyday, such as potatoes, apples, sweet potatoes and strawberries
are polyploid.
An interesting example is bananas. Bananas are triploid, and typically
triploid organisms do not make the sperm or eggs needed to reproduce.
Thus, bananas are effectively sterile. That means you can't get banana
seeds to make more bananas. Farmers cut off shoots from the side of the
banana plant before it produces fruit and dies, and propagate them
asexually.

However, polyploidy is especially common among ferns and flowering

plants, including both wild and cultivated species. Wheat, for example, after
millennia of hybridization and modification by humans, has strains that are
diploid (two sets of chromosomes); tetraploid (four sets of chromosomes),
with the common name of durum or macaroni wheat; and hexaploid (six
sets of chromosomes), with the common name of bread wheat.
Many agriculturally important plants of the genus Brassica are also
tetraploids. This genus, known as cabbages or mustards, includes turnips,
brussels sprouts, cabbage, cauliflower, broccoli, mustard seed and other
important crops. The Triangle of U is a theory, developed by a Woo Jang-
choon, a Korean botanist who was working in Japan, that says the
genomes of three ancestral species of Brassica combined to create the
three common tetraploid species Brassica juncea (Indian mustard),
Brassica napus (Rapeseed, rutabaga), and Brassica carinata (Ethiopian
mustard).

However, polyploidy is especially common among ferns and flowering

plants, including both wild and cultivated species. Wheat, for example, after
millennia of hybridization and modification by humans, has strains that are
diploid (two sets of chromosomes); tetraploid (four sets of chromosomes),
with the common name of durum or macaroni wheat; and hexaploid (six
sets of chromosomes), with the common name of bread wheat.

Many agriculturally important plants of the genus Brassica are also

tetraploids. This genus, known as cabbages or mustards, includes turnips,
brussels sprouts, cabbage, cauliflower, broccoli, mustard seed and other
important crops. The Triangle of U is a theory, developed by a Woo Jang-
choon, a Korean botanist who was working in Japan, that says the
genomes of three ancestral species of Brassica combined to create the
three common tetraploid species Brassica juncea (Indian mustard),
Brassica napus (Rapeseed, rutabaga), and Brassica carinata (Ethiopian
mustard).
In plant breeding, the induction of polyploids is a common technique to
overcome the sterility of a hybrid species. Triticale is the hybrid of wheat
(Triticum turgidum) and rye (Secale cereale). It combines sought-after
characteristics of the parents, but the initial hybrids are sterile. After
polyploidization, the hybrid becomes fertile and can thus be further
propagated to become triticale.
Polyploid plants in general are more robust and sturdy than diploids. In the
breeding of crops, those plants that are stronger and tougher are selected.
Thus, many crops have unintentionally been bred to a higher level of
ploidy:
Triploid crops: banana, apple, ginger
Tetraploid crops: durum or macaroni wheat, maize, cotton, potato,
cabbage, leek, tobacco, peanut, kinnow, Pelargonium
Hexaploid crops: chrysanthemum, bread wheat, triticale, oat
Octaploid crops: strawberry, dahlia, pansies, sugar cane

Polyploidy in fungi:
Fungi seems to stand in a zone intermediate between prokaryotes and the
multicellular organism and plants with respect to their plant sizes and
complexity “if indeed the fungi posses the limited evolutionary potential
because of limited genetic material, any means to increase the quantity of
DNA should provide a mean to escape this potential. The best known
mechanism for this should be polypolidization.
Polyploidy in fungi has been considerd to be non-existance or rare. And
this assumption would indicate that polyploidy as an evolutionary
mechanism for fungal speciation has not assumed the same sort of role
accorded it in the evolution of vascular plant.
Several example of polyplid are:
autopolyploid: the aquatic fungi of genus Allomyces some
Saccharomyces cerevisiae strains used in bakery
allopolyploid: the widespread Cyathus stercoreusthe allotetraploid lager
yeast Saccharomyces pastorianus,[43] the allotriploid wine spoilage yeast
Dekkera bruxellensis
paleopolyploid: the human pathogen Rhizopus oryzae,

As for plants and animals, fungal hybrids and polyploids display structural
and functional modifications compared to their progenitors and diploid
counterparts. In particular, the structural and functional outcomes of
polyploid Saccharomyces genomes strikingly reflect the evolutionary fate of
plant polyploid ones. Large chromosomal rearrangements leading to
chimeric chromosomes have been described, as well as more punctual
genetic modifications such as gene loss The homoealleles of the
allotetraploid yeast S. pastorianus show unequal contribution to the
transcriptome Phenotypic diversification is also observed following
polyploidization and/or hybridization in fungi producing the fuel for natural
selection and subsequent adaptation and speciation.

References:
https://masters.agron.iastate.edu/classes/527/lesson09/09.02.html
http://www.biology-pages.info/E/Endoreplication.html#Polyploidy
https://en.wikipedia.org/wiki/Polyploid#Types
http://medicine.jrank.org/pages/2667/Polyploidy-Polyploidy-in-Animals.html
http://study.com/academy/lesson/polyploidy-in-plants-animals-speciation-
cells.html
http://www.newworldencyclopedia.org/entry/Polyploidy
http://link.springer.com/chapter/10.1007%2F978-1-4613-3069-1_9#page-2