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360 CELL BIOLOGY

C H A P T E R

22
Parthenogenesis

U
sually an unfertilized ovum develops into a new
individual only after the fertilization but in certain
cases the development of the egg takes place without
the fertilization. This peculiar mode of sexual reproduction in
which egg development occurs without the fertilization is
known as the parthenogenesis (Gr., parthenos=virgin;
genesis=origin). An organism that has developed parthenoge-
netically is called a parthenogenone or parthenote. The
phenomenon of the parthenogenesis occurs in different groups
of the animals as in certain insects (Hymenoptera, Homoptera,
Coleoptera), crustaceans, rotifers and also in some vertebrates
such as several desert lizards, turkeys and some mammals.
There are certain conditions which are intermediate
between parthenogenesis and fertilization, e.g., partial fertili- In spring and early summer, when food
zation, gynogenesis, androgenesis and merogony. 1. In par- is abundant, aphid females reproduce
tial parthenogenesis, the egg may be fertilized by only a part parthenogenetically.
of sperm. For example, according to Boveri in sea urchin egg
the fertilization of the egg (activation) takes place by the
sperm aster. The sperm nucleus gets fused with the egg
nucleus only in two cell stage. 2. In gynogenesis, the sperm
penetrates the egg but takes no part in development. It degen-
erates in the egg without fusion with the egg nucleus, e.g.,
Rhabditis aberrans. 3. In androgenesis, the egg is activated
by the sperm and development takes place without the partici-
pation of the egg nucleus. For example, if the ova of frogs and
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PARTHENOGENESIS 361
toads are treated with radium and then fertilized by normal sperms, the egg nucleus does not take part
in development, but sperm (paternal) nucleus participates in normal development. 4. In merogony,
egg fragments devoid of nucleus develop when fertilized by a normal sperm. If sea urchin eggs are
shaken to produce small pieces, the fragments round up to form spheres. Some of these spheres are
without nuclei. If such enucleated spheres are normally fertilized, they may develop into dwarf larvae.
The parthenogenesis may be of two types :
1. Natural parthenogenesis; 2. Artificial parthenogenesis.
NATURAL PARTHENOGENESIS
In certain animals the parthenogenesis occurs regularly, constantly and naturally in their life
cycles and is known as the natural parthenogensis. The natural pathenogenesis may be of two types,
viz., complete or incomplete.
(i) Complete parthenogenesis. Certain insects have no sexual phase and no males. They
depend exclusively on the parthenogenesis for the self-reproduction. This type of parthenogenesis is
known as the complete parthenogenesis or obligatory parthenogenesis. It is found in some species
of earthworms, badelloid rotifers, grasshoppers, roaches, phasmids, moths, gall flies, fishes, sala-
manders and lizards.
(ii) Incomplete parthenogensis. The life cycle
of certain insects includes two generations, the sexual
generation and parthenogenetic generation, both of
which alternate to each other. In such cases, the
diploid eggs produce females and the unfertilized
eggs produce males. This type of parthenogenesis is pg 100
genetics & evolution
known as the partial or incomplete or cyclic par-
thenogenesis. book with Shubha
Cyclic parthenogenesis shows several varia-
tions in the alternation of sexual (S) and parthenoge-
netic (P) generations : (1) In gall flies (e.g.,
Neuroterus) there is an alternation of one sexual and
one parthenogenetic generation per year (P, S,....P,
S,.... P, S). (2) In aphids (plant lice), daphnids and
rotifers, the sexual generation may come after many In the summer water fleas reproduce very rapidly by
parthenogenesis. 10-12 unfertilized eggs are seen
parthenogenetic generations during the summer of here inside the brood pouch of female.
the year (P, P, P, P, P, P, S,.... P, P, P, P, P, P, S....).
(3) In gall midge (Miaster) the larvae reproduce indefinitely by paedogenetic parthenogenesis. In
this case, germ cells within the larvae develop parthenogenetically into parasitic larvae which feed on
the mother larvae. These larvae tend to live under the bark of rotting logs and feed on fungus. Under
favourable conditions winged males and females are produced. These stages reproduce sexually and
help in dispersion. (4) In some groups there is no regularity between parthenogenetic and sexual
generations.
The complete and incomplete type of natural parthenogenesis may be of following two types :
(a) Haploid or arrhenotokous parthenogenesis; (b) Diploid or thelytokous parthenogenesis.
(a) Haploid or arrhenotokous parthenogenesis. In the arrhenotokous parthenogenesis, the
haploid eggs are not fertilized by the sperms and develop into the haploid individuals (Fig. 22.1). In
these cases the haploid individuals are always males and the diploid individuals are the females e.g.,
1. Insects : (i) Hymenoptera (Bees and Wasps) (ii) Homoptera, (iii) Coleoptera (Micromalthus
debilis), (iv) Thysanoptera (Anthothrips verbasi). 2. Arachnids, e.g., ticks, mites and certain spiders
(Pediculoids ventricusm). 3. Rotifers, e.g., Asplanchne amphora.
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362 CELL BIOLOGY

Thus, the queen bee is fertilized only once by one or many males (drones). She stores the sperm
in her seminal receptacles and as she lays her eggs, she can either fertilize the eggs or allow them to
pass unfertilized. The fertilized eggs become females (fertile queens or sterile workers depending up
on the amount of royal jelly the developing young receives); the unfertilized eggs become fertile males
or drones.

2N
queen bee

meiosis

N mature ova

sperm stored
in sperm sac
of queen

2N zygote N zygote (haploid)

m
ito
sis

2N female worker N haploid male

bee

2N 2N N
2N
N N N
2N 4N 2N N
N activation
oogonium tetrad 1st polar body 2nd meiotic 2nd polar 1st cleavage
metaphase body
Fig. 22.1. Schematic representation of haploid parthenogenesis in bees (after Grant, 1978).

(b) Diploid or thelytokous parthenogenesis. In the diploid parthenogenesis, the young

individuals develop from the unfertilized diploid eggs. The offspring of thelytoky could theoretically
be either male or female; but normally it produces only diploid females (Fig. 22.2). For example, in
aphids, females emerging in the spring produce several generations of females by diploid partheno-
genesis resulting from suppression of first or second polar body. At summer’s end some females
produce sexual males and females by diploid parthenogenesis, males differing from females in lacking
one sex chromosome. Males produce haploid gametes through normal meiosis which fuse to form
diploid zygotes that emerge again in the spring as parthenogenetic females. Further, since thelytoky
is also found in polyploid forms, it is also called somatic parthenogenesis.
Following types of the thelytoky have been recognised :
(i) Ameiotic parthenogenesis. Sometimes during the oogenesis, first meiotic or reduction
divsion does not occur but second meiotic division occurs as usual. Such eggs contain diploid number
of chromosomes and develop into new individuals without the fertilization. This type of parthenogen-
esis is known as apomiotic or ameiotic parthenogenesis and occurs in Trichoniscus (Isopoda),
Daphnia pulex (Crustacea) Compelona rufum (Mollusca), weevils and long-horned grasshopper.
(ii) Meiotic parthenogenesis. Certain eggs develop by the usual process of oogenesis but at
certain stages diplosis or doubling of chromosome number and production of diploid eggs occur. Such
eggs develop into the diploid individuals and this phenomenon is known as the meiotic parthenogenesis.
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PARTHENOGENESIS 363

2N
zygote 2N
gametes
N
2N
spring summer
partheno-
genetic
females
meiosis

2N
2N – 1 2N – 1
adult male
2N
stem
mother

2N female sexual 2N ova male and female

forms (partheno - genetic) producers

2N 2N 2N
activation
N
2N 4N 2N 2N 2N 2N
N
oogonium tetrad 1st polar body 2nd meiotic suppression of 1st cleavage
metaphase 2nd polar body
Fig. 22.2. Schematic representation of diploid parthenogenesis in aphids (after Grant, 1978).

The diplosis of the diploid thelytoky may occur by the following methods :
(a) By autofertilization. In certain cases the oocyte divides meiotically up to the formation of
ootid and secondry polocyte. But the ootid and the secondary polocyte unite together to form a diploid
egg which develops into a new individual, e.g., Artemia salina (Crustacea) and various other
organisms.
(b) By restitution. Sometimes in primary oocyte karyokinesis forms a nucleus of the secondary
oocyte and a nucleus of the first polocyte. But the karyokinesis is not followed by the cytokinesis. The
chromosomes of both daughter nuclei are arranged on the equator and undergo second meiotic division
to form a diploid ootid and a diploid polocyte. The dipioid ootid or ovum develops into a
parthenogenetic diploid individual. This type of diplosis is known as the restitution, e.g., insects of
order Hymenoptera (Nemertis conesceus) and Lepidoptera.
Natural Parthenogenesis in Vertebrates
A few cases of natural parthenogenesis have also been reported in the vertebrates. The fish
Carassius auratus gibelio is reported to consist of females only (Lieder, 1955). Likewise, males are
found totally lacking in the lizard Lacerta sexicola armeniaca (Lantz and Cyren, 1936). In it females
are reported to be originated by parthenogenesis. In turkeys 80 per cent of incubated eggs show early
cleavage stages. Such parthenogentic forms have hatched and grown to reproducing adults which are
found to be diploid male with ZZ sex chromosomes. In mammals too, up to 60 per cent of hamster eggs
becomes spontaneously activated and develops up to two-cell stage (Austin, 1956).
ARTIFICIAL PARTHENOGENESIS
The eggs which always develop into the young individuals by the fertilization sometimes may
develop parthenogenetically under certain artificial conditions. This type of parthenogenesis is known
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364 CELL BIOLOGY

as artificial parthenogenesis. The artificial parthenogenesis may be induced by various physical and
chemical means.
A. Physical means. The following
physical means cause the parthenogen-
esis :
(i) Temperature. The range of tem-
perature may induce parthenogenesis in
the eggs. For instance, when the egg is
transferred from the 30o C to 0–10o C the
parthenogenesis is induced.
(ii) Electrical shocks can cause
parthenogenesis.
(iii) Ultraviolet light can cause par-
thenogenesis.
(iv) When the eggs are pricked by
the fine glass needles the development of This whiptail lizard reproduces by
young ones takes place parthenogeneti- parthenogenesis.
cally.
B. Chemical means. The following chemicals have been found to cause parthenogenesis in the
normal eggs :
1. Chloroform 2. Strychnine 3. Hypertonic and Hypotonic sea waters. 4. Chlorides of K+, Ca++,
Na , Mg++, etc. 5. Acids as butyric acid, lactic acid, oleic acid and other fatty acids. 6. Fat solvents,
++

e.g., toulene, ether, alcohol, benzene and acetone. 7. Urea and sucrose.
The artificial parthenogenesis has been induced by above mentioned physical and chemical
means by various workers in the eggs of most echinoderms, molluscs, annelids, amphibians, birds and
mammals.
SIGNIFICANCE OF PARTHENOGENESIS
1. The parthenogenesis serves as the means for the determination of sex in the honey bees, wasps,
etc., and it supports the chromosome theory of inheritance.
2. The parthenogenesis is the most simple, stable and easy process of the reproduction, e.g.,
aphids (insects).
3. The parthenogenesis eliminates the variation from the population, but encourages develop-
ment of the advantageous mutant characters.
4. The parthenogenesis causes the polyploidy in the organisms.
5. Due to the parthenogenesis, there is no need for the organisms to waste their energy in the
process of mating but it allows them to utilize that amount of energy in the feeding and reproduction.
6. Honey bee and other social insects also control their sex ratio by parthenogenesis.
7. In aphids, parthenogenesis is a means of rapid breeding; the females reproduce by diploid
parthenogenesis during summer.
REVISION QUESTIONS
1. What is parthenogenesis ? Describe different types of natural parthenogenesis in animals.
2. Give an account of artificial parthenogenesis.
3. What is the significance of parthenogenesis ?