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Cytoplasmic Inheritance
(With Diagram) | Cell
Biology
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In this article we will discuss about Cytoplasmic

Inheritance:- 1. Subject-Matter of Cytoplasmic
Inheritance 2. Terms and Definitions
of Cytoplasmic Inheritance 3. Characteristics and
Detection 4. Extra-Nuclear Inheritance in
Eukaryotes.
Subject-Matter of Cytoplasmic Inheritance:
The existence of genes as segments of nucleic acid
molecules, located in chromosome of nucleus, has been
demonstrated by several experiments. The nuclear genes
control the phenotypes of the organisms and are concerned
with the transmission of hereditary character from one
generation to next generation is known and predictable
Mendelian fashion.
The inheritance of genes of nuclear chromosomes is
characterised by the fact that the genes from male and
female parents contribute equally to the genetic
constitution of the offspring. Therefore, in it the reciprocal
crosses between parents of different homozygous genotype
will produce offspring’s of identical phenotypes except for
sex-linked genes.
However, in certain cases, although male and female
parents contribute equally their nuclear genes to the
offspring’s, the results show a non-Mendelian inheritance
pattern and the result of reciprocal crosses varies.

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These variations suggest that the genes for the inheritance

of certain characters do not occur within the nucleus, but
they are present ill cytoplasm and play an important role in
transmission of certain specific traits, which are not
controlled by nuclear genes. Therefore, it builds up the
concept of cytoplasmic inheritance. The genes for
cytoplasmic inheritance are independent, self-replicating
nucleic acids.
Evidence for cytoplasmic inheritance was first reported by
Correns in Mirabilis jalapa and by Bar in Pelargonium
zonule in 1908. Rhoades described cytoplasmic male
sterility in maize in 1933. In 1943, Sonneborn discovered
kappa particles in Paramoecium and described its
cytoplasmic inheritance. Presence of DNA in chloroplasts
was first demonstrated by Ris in plant cell.
In 1963, Nass and his co-workers proved the existence of
DNA in mitochondria. Subsequently, from time to time,
observations by several scientists have been reported the
important role of cytoplasm in genetics. Thus, on the basis
of observations made on cytoplasmic inheritance of some
specific traits, it has been suggested that cytoplasm is also
genetically active.

Terms and Definitions of Cytoplasmic

Inheritance:
Extra-chromosomal inheritance, extra-nuclear inheritance,
somal inheritance and maternal inheritance are all
synonyms. All these terms can be defined as the inheritance
of characteristics of only one of the two parents, usually the
female parent to the progeny. The reciprocal crosses show
consistent differences as well as there is a lack of
segregation in F2 and subsequent generations.
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The genes controlling cytoplasmic inheritance are present

outside the nucleus and, in the cytoplasm, they are known
as plasma genes, cytoplasmic genes, cytogeneses, extra
nuclear genes or extra chromosomal genes.
The sum total of the genes present in cytoplasm of a cell is
known as Plasmon. All the genes present in a plastid are
known as plastoms. Similarly, all the genes present in a
mitochondrion are known as chondrioms. The genes
present in plastid and in mitochondrion are located in their
own DNAs and are known as cp DNA and mt DNA,
respectively. These DNAs are collectively termed organelle
DNA.

Characteristics and Detection of Cytoplasmic

Inheritance:
Cytoplasmic inheritances do not show Mendelian
inheritance.
They show the following characteristic features:
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i. Hereditary traits which are transmitted by cytoplasm do

not show Mendelian segregation in crosses and in
reciprocal crosses with respect to a particular set of charac-
teristics controlled by a set of cytoplasmic genes produce
dissimilar hybrids.
ii. Most of the recorded cytoplasmically inherited
characteristics would follow the maternal line, i.e.,
uniparental mode of transmission. In higher plants and ani-
mals, ovum or egg cell is comparatively large and contains
large amount of cytoplasm. But male gametes or sperms
have very little amount of cytoplasm. So, under this
situation, most of cytoplasmic factors are transmitted to the
progeny through the ovum of mother.
It is known as maternal inheritance or trans-ovarian trans-
mission. In this mode of transmission, all the offspring’s of
the parents have maternal condition and only female
progeny can transmit the cytoplasmic characteristics to the
succeeding generations. Hence the reciprocal crosses yield
different or non-Mendelian results.
Characteristics of Mendelian Inheritance:
The inheritance pattern of characters of an organism as
proposed by Mendel on the basis of monohybrid and di-
hybrid crosses is referred to as Mendelian inheritance.

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It shows the following characteristic features:

i. Contribution of both male and female is equal, hence
results from reciprocal crosses are similar.
ii. Segregation produces the phenotypes ratio 3 : 1 and
genotype ratio 1 : 2 : 1 in the F2 generation of a monohybrid
cross and a typical phenotype ratio 9 : 3 : 3 : 1 in di-hybrid
crosses.
Mendelian inheritance pattern is regarded as a sufficient
evidence for a gene to be located in chromosomes; such
genes are called nuclear genes or simply as genes.

Extra-Nuclear Inheritance in Eukaryotes:

Various cases of extra-nuclear inheritance in different
eukaryotic organisms have been studied by several
scientists.
Few important examples of extra nuclear
inheritance in eukaryotes are stated under some
classified subheadings:
(i) Maternal Inheritance:
Maternal inheritance means the inheritance controlled by
extra-chromosomal, i.e., cytoplasmic, factors that are
transmitted to the succeeding generation through the egg of
female organism.
They show the following features:
i. reciprocal differences in F1;
ii. which in most cases disappears in F2;
iii. a smaller variation in F2 as compared to that in F3.
Maternal inheritance may be, broadly speaking, of
two kinds:
i. If some treatments (chemical poison, heat shock etc.) are
applied to the female parent, it may affect the egg’s
cytoplasm. As a result subsequent offspring’s are modified
in some way. Effects of this kind are called Dauer-
modifications or persisting modifications.
It is observed that when protozoa are treated
experimentally with chemical poisons or heat shocks, the
treatments induce several morphological abnormalities in
them. Such abnormalities go on decreasing generation after
generation and, eventually, disappear completely through
cell division if the treatments are removed.
Further evidences also come from fruit flies subjected to
heat treatment and from bacteria treated with chemicals.
ii. Other kinds of maternal inheritance are also known
which do not depend upon the repeated application of an
external stimulus to the cytoplasm. In this case, maternal
inheritance is truly controlled by independent cytoplasmic
genes.
Maternal effects reflect the influence of the mother’s gene
on developing tissues. Many important characteristics of
both animal and plants show maternal effects of which
some examples axe described next.
(ii) Coiling of Snail Shells (Limnaea Peregra):
One of the earliest and classical examples of a maternal
effect is that of the direction of coiling in shells of the water
snail Limnaea peregra. In this snail, the shell is spirally
coiled. Usually the direction of coiling of the shell is clock-
wise if viewed from the top of the shell. This type of coiling
is called dextral. However, in some snails the coiling of shell
is anticlockwise. This type of coiling is sinistral.
The direction of shell coiling of both types of snail is
governed by genotype of the female parent and not by their
own genotype. Further investigation suggests that coiling
depends upon the early clearage in the zygote.
If the mitotic spindle is tilted to left (Fig. 22.1) of the
median line of zygote, the successive cleavages will produce
a spiral to left (sinistral) and if the orientation of spindle is
tilted to the right of the median line of zygote, the
successive cleavages will produce a spiral to right (dextral).
The spindle orientation is controlled by the genotype of
oocyte from which the egg develops.
In a cross (Fig. 22.2) between dextral 9 (female) snail and
sinistral (male), (follow the left side coloum of the Fig. 22.2)
all the Fi progeny have dextral coils like their mother and
also indicates that dextral character (RR) is dominant over
sinistral coiling (rr).
However, in the F1 x F1 cross (i.e., inbreeding or self
fertilisation) all the F2 snails are also dextral. The F3 progeny
from F2 individuals with the genotype RR and Rr will show
dextral coiling while those from rr F2 individual will exhibit
sinistral coiling of their shell; this produces the typical 3 : 1
ratio in F3 generation.
In reciprocal cross (right side column of the Fig. 22.2)
between dextral (male) and sinistral (female), all the
Fx progeny have sinistral coiling (Rr) instead of dextral
coiling. In this case, F1 x F1 cross, all the F2 snails are, again,
dextral. This F3 progenies from F2 also exhibit the typical
monohybrid ratio of 3 : 1.
Thus the features of inheritance of coiling of
Limnaea shells may be summarised as:
i. F1s from reciprocal crosses show differences in coiling
pattern.
ii. No segregation in F2.
iii. Appearance of the typical 3: 1 ratio in F3 in place of F2.
The 3 : 1 ratio, although in F3, clearly indicates that coiling
of shell is governed by a single nuclear gene. But the
segregation of this nuclear gene is apparently delayed by
one generation and is visible in F3instead of F2 as in all other
cases of Mendelian inheritance. This is because the
direction of coiling in this snail is primarily determined by
some substances already present in the cytoplasm or
ooplasm of the egg cell.
Obviously these substances are produced by the female
parent. As a result the offspring would produce the
phenotype (in F1) of the maternal parent since its nuclear
gene product is possibly active after one generation later
and shows delayed segregation in F3.
(iii) Maternal Inheritance in Drosophila:
Several examples of maternal effects are known in
drosophila:
i. Abnormal growth in the head region of Drosophila
melanogaster was produced sporadically in a sample from a
wild population collected at Acahuizotla, Mexico.
Development of abnormal growth in the head region is
called Tumorous head (Tu – h). Tu – h is governed by two
major genes. But the frequency of tumour development in
progeny is markedly influenced by the maternal effect.
When a cross is made between a normal female fly and a
male fly with head tumour, less than 1% of the progeny
exhibit head tumour. In contrast, when a reciprocal cross is
made between a female having head tumour and a normal
male, about 30% of the progeny show tumour development.
ii. In Drosophila, fertility and survival are occasionally
influenced by maternal effect. A recessive nuclear gene,
grandchild-less, affects the fertility of progeny in Drosophila
subobscura. In this fly a female homozygous for which is
fertile but all her offspring’s are sterile.
The reason of this effect arises from cytoplasmic dependent
pathway for the development of many organs, the egg
cytoplasm formed by a female fly is not uniform and various
parts of the egg appear to be specifically assigned for the
formation of different tissues. Thus the fate of Drosophila
germ cells to produce either various or testes is determined
early in development.
A daughterless (da) gene in Drosophila causes death of all
XX zygotes derived from eggs of da da females. It is
reported that the cytoplasm of eggs of da da females affects
the two X-chromosomes and does not inhibit the
development of the female phenotype.
It should be noted that the action of da is determined by the
genotype of the female producing egg and not by the
genotype of the egg cells themselves. It is also reported that
interaction of da gene product is evident if the zygote is XX
but there is no such action affecting the survival of XY
zygote.
Some genes in Drosophila have lethal effects on male
embryos. A recessive gene, sonless, leads to the absence of
males in the progeny of females homozygous for this gene.
Another gene, abnormal oocyte, reduces the frequency of
males in the progeny.
These effects on the frequency of male progeny are
produced due to effect of egg cytoplasm on the survival of
XY zygote. Thus the genes for daughterless and sonless
produce typical uniparental inheritance. These uniparental
inheritance patterns depend on the nuclear genotype of the
female parent.
(iv) Maternal Inheritance in Ambystoma:
Maternal inheritance has also been studied in Axololt of
Ambystoma maxicanum. In this animal there is a lethal
gene ‘O’. It is a recessive gene. But its dominant gene is ‘+’.
The heterozygous individuals with genotype ‘+O’ are
completely normal.
When two heterozygous individuals with genotype ‘+0′ are
crossed, a homozygous offspring with genotype ’00’ may be
produced. The homozygous individual (‘OO’) develop
normally in the early stage but in later life show a slight
retardation of growth.
Their regeneration capacity is reduced to great extent and
the homozygous males (OO) are sterile and their testes are
poorly developed and spermatogenesis does not produce
beyond spermatogonial stages. On the other hand, the
homozygous females with genotype ‘OO’ produce eggs.
The eggs are capable of fertilisation and normal clearage but
at the onset of gastrulation, further development is retarded
and embryos normally die. This abnormal course of
development is affected by the maternal effect, i.e., the
growth is resulted by the genotype of the mother
irrespective to that of the offspring.
If the maternal genes are recessive, i.e., ‘OO’, the growth of
the offspring even with genotypes ‘+O’ is retarded.
According to Biggs and Justus (1967), in the egg of
homozygous female ‘OO’ a protein-like substance called
corrective factor is absent.
This protein-like substance is necessary for normal growth
and development of the offspring and is synthesised by
normal genes ‘++’ of homozygous or heterozygous normal
females (‘++’ or ‘+O’) during oogenesis.
(v) Maternal Inheritance of Eye Pigmentation in
Water Flea and Flour Moth:
The maternal inheritance has also been observed in case of
water flea (Gammarus sp.) and the flour moth (Ephestia
kuhniella). The normal colour of both the invertebrates is
dark due to the dominant gene (AA or KK) in which the
dominant gene A or K directs the production of a hormone-
like substance called Kynurenine which is involved in the
pigment synthesis.
The recessive gene (i.e., a or k) fails to direct the synthesis
of kynurenine. In absence of kynurenine, colour of eye
becomes light.
Therefore the recessive mutants do not possess pigment in
the eye and have the genotype aa or Kk. When a cross is
made between a heterozygous male (Aa or Kk) and a double
recessive female (aa or kk), only half of the larvae show dark
pigment in the eye. Again, when a cross is made between a
heterozygous female and a double recessive male, all larva
are with dark eye. But on reaching the adult stage, half of
the progenies having the genotype ‘aa’ or ‘kk’ become light-
eyed.
This indicates that some kynurenine molecules diffuse from
the ‘Aa’ mother into all young (larvae) enabling them to
produce pigments regardless of their genotype. But the aa
progeny is unable to synthesize kynurenine and,
consequently, develops light eye as the kynurenine obtained
from mother is used up. This example suggests an
ephemeral type of maternal inheritance.
(vi) Maternal Inheritance in Mammals:
Maternal effects in mammals are very prominent because
mammalian mothers may affect the development of their
offspring’s not only through ovum cytoplasm but also
through the uterine environment. Effects of the maternal
Rh blood group on the developing foetus in humans and
that of maternal genotype for hair loss gene on the survival
of young embryos in mice are the examples of maternal
effect.
Other examples of maternal effects in human are: the
embryonic defects caused by maternal diabetes and
maternal phenylketonuria, a maternal influence on left-or
right-handedness and a maternal effect on body weight.
(vii) Maternal Inheritance in Plants:
Cytoplasmic male sterility in cross-pollinating plants is a
classic example of maternal inheritance. Plants which are
unable to produce functional pollen but possess normal
fertile female structure aired known as male sterile.
Thus male sterility is characterised by nonfunctional pollen
grains. This occurs in many flowering plants. In maize,
wheat, sugar-beets, onions and some other crop plants,
fertility is controlled, at least in part—by cytoplasmic factors
and results in male sterility.
This type of male sterility is referred to as cytoplasmic male
sterility. Nuclear gene did not control this type of sterility,
rather it is transmitted from generation to generation
through egg cytoplasm. In other plants, however, male
sterility is controlled entirely by nuclear genes. But over
present discussion will be focused on only cytoplasmic male
sterility in relation to maternal inheritance in plants.
Cytoplasmic male sterility is determined by cytoplasmic
factors. Since the bulk of cytoplasm of zygote is contributed
by the egg cell and the pollen tube containing male gametes
contributes negligible or no cytoplasm, sterility factors
present in the cytoplasm of egg cell will be transmitted to
the offspring which would always be male sterile.
A case of cytoplasmic male sterility was discovered and
carefully analysed by M. M. Rhoades (1933) in maize. He
observed a male sterile plant in maize. In this plant pollens
are aborted in the anther. The male sterile plant is produced
when an egg cell containing cytoplasmic male sterility factor
is fertilised by pollen from normal male fertile plants.
It indicates that male sterility is contributed by the
cytoplasm of female parent. It is also- observed that when a
male sterile female plant is crossed with wide range of
fertile males, all progenies axe male sterile in the
subsequent generations.
In maize, three distinct male sterile source (Cms)
cytoplasm’s axe known which are designated as Cms-T,
Cms-C and Cms-S. The normal male fertile cytoplasm is
known as N- cytoplasm. Each of the three Cms cytoplasm’s
shows strict maternal inheritance—even when all
chromosomes axe replaced from male sterile plants by a
male fertile source through repeated backcrossing.
Even then male sterility characteristics could not be avoided
and the characteristics till persists. It indicates that if the
character is present on chromosome as nuclear gene, then
male sterility could be eliminated by repeated backcrossing.
Therefore, it is confirmed that male sterility is not
controlled by nuclear gene, i.e., nuclear gene has no
influence on cytoplasmic male sterility. In rare cases, male
sterile plants produce a few fertile pollen grains.
When reciprocal crosses axe made between male sterile
parent (with fertile pollen) and normal male fertile
(female), the progeny is found to be male fertile. Such cases
confirm maternal inheritance of male sterility.
Recent studies have shown that cytoplasmic male sterility is
controlled by either some unique polypeptide produced by
mitochondria or by some plasmid-like elements which axe
not found in the mitochondria of normal cytoplasm.
Cytoplasmic male sterility due to S-cytoplasm is different
from the cytoplasmic male sterility due to T-cytoplasm in
several ways. The mt DNA of S-cytoplasm contains two
unique plasmid-like DNA fragments called SF (Mol. wt.
3.45 x 106) and S-S (mol.wt.4.10 x 106).
These plasmid-like DNAs are not found in the isolated DNA
of chloroplast or nuclei. Therefore, these plasmid-like DNAs
are the characteristic of mitochondrial DNA of S-cytoplasm.
These plasmid-like DNAs are also absent in the
mitochondrial DNA of N- cytoplasm (Normal-fertile), T-
cytoplasm (male sterile) as well as C-cytoplasm (male
sterile). Hence it has been suggested that such plasmid- like
DNAs in mitochondria are responsible for causing male
sterility.
T and C-cytoplasm of male sterile plant is comparatively
stable and irreversible. It means that they never give rise to
fertile cytoplasm even by applying mutagens.
On the other hand, S-cytoplasm is stable. It is found
to change into fertile condition in some cases due
to one of the two kinds of change:
i. Cytoplasmic mutation from male sterile to male fertile;
ii. Nuclear mutation giving rise to a new repressor gene.
It is also shown that both these changes are involved to
make it fertile.
When a fertile line derived from S-cytoplasm is crossed as
male ( ) with a Cm-S tester female plant ( ), in some
cases the offspring is male sterile. It indicates that the
restorer gene is, possibly, absent. In other cases the
offspring is semi-fertile. It suggests that the restorer gene is
possibly, present. This restorer gene is different from the
normal nuclear restorer gene – Rf3 meant for S-cytoplasm.
These new restorer genes are likely to be located on
different chromosomes where they are, possibly, attached
like episome at different times and bring the change from
sterile to fertile condition. It has also been suggested that
male fertility genes could be originally located on organelle
DNA and were later transposed to a nuclear site giving rise
to restorer gene.
The gene or DNA segment that has migrated from the
organelle to the nucleus or to the other organelle is termed
promiscuous DNA.
When this fertility gene is absent from both organelle and
nucleus, this might have led to cytoplasmic male sterility.
The restorer gene present in the nucleus as dominant gene
generally nullifies the effect of cytoplasmic male sterility so
that individuals having a restorer gene in homozygous or
heterozygous state are fertile even in the presence of male
sterile cytoplasm.
In case of Cms-T, plasmid-like event in the mitochondria is
absent but some unique polypeptides are produced in the
mitochondria which bring the male sterility. When restorer
gene is present in the nucleus, it prevents the production of
unique polypeptides in Cms-T, and the plant becomes
fertile.
But when nuclear restorer gene is absent, the plant achieves
male sterile cytoplasm. Cms-C has also two additional
plasmid-like elements like Cms-S. These elements are
associated with cytoplasmic male sterility.
The maternal inheritance mechanism that transmits male
sterility in maize have also been demonstrated by Dhawan
and Paliwal in 1964. In their experiment they used two
strains of maize—Sikkim primitive-2 and another strain
from Colorado—for reciprocal crosses.
When Sikkim primitive-2 was used as female parent in the
cross, the offspring showed little vigour and poor yield, but
when Colorado strain of maize was used as female parent,
the hybrid were more vigorous and showed high yield
potency. These differences in hybrids of reciprocal crosses
suggests that yield and vigour are governed by female
cytoplasm.
Extra-nuclear Inheritance by Cellular Organelles:
Extra-nuclear inheritance is also associated with certain
cytoplasmic organelles (mitochondria, plastids) that contain
naked circular DNA and protein synthesizing apparatus.
These extra nuclear genetic materials present in the
organelles axe autonomous and code only for limited
number of enzymes and polypeptides. Certain enzymes
required for cellular respiration are synthesised in the
mitochondria.
Similarly, chlorophyll and other pigments axe synthesised
in the plastid. Besides the involvement of such biosynthetic
activities, these organelles DNAs axe directly associated
with the inheritance of some phenotypes which are not
controlled by the nuclear genes. The genetic material of
chloroplasts and mitochondria are transmitted almost
exclusively via the egg.
The inheritance pattern is well-illustrated by the
following examples:
Plastid Inheritance in Mirabilis:
Plastid inheritance means the inheritance of plastid
characteristics due to plasma genes located in plastids.
Plastid inheritance was first described by C. Corens (1908)
in the four o’clock plant, Mirabilis jalapa.
Leaves of Mirabilis jalapa may be green, white or variegated
and some branches may have only green, only white or only
variegated leaves. Variegation means the presence of white
or yellow spots of variable size on the green background of
leaves.
Thus it forms the mosaic pattern of coloration on a leaf. Due
to certain inheritable defects chloroplast of all cells or some
cells of leaf often are unable to synthesize the chlorophyll
pigments. Such cells remain non- green and form white or
yellow coloured leaf, or white or yellow patches,
interspersed with areas containing normal green cells with
healthy chloroplasts.
Variegation may be produced by:
(a) Some environmental factors,
(b) Some nuclear genes,
(c) Plasma-genes in some cases.
Since the first and second causes of leaf variegation do not
concern cytoplasmic inheritance, the inheritance of
variegation due to plasma-genes will be discussed in this
article.
Correns made reciprocal crosses (Fig. 22.3) in all
combinations among the flowers produced on these three
types of branches.
These results are summarised in Table 22.1.:

The results obtained from various crosses of leaf

phenotypes of Mirabilis jalapa, as shown in Table 22.1,
clearly indicates that leaf phenotype of the progeny is the
same as that of the female parent (Fig. 22.3). The phenotype
of male parent did not contribute anything to the progeny.
This phenomenon is referred to as uniparental
transmission. Again, the results of the crosses of Mirabilis
jalapa cannot be explained by sex-linkage.
The inheritance of different leaf colours in Mirabilis jalapa
might be explained if the plastids are somehow autonomous
and are never transmitted through male parent. For an
organelle to be genetically autonomous, it must be provided
with its own genetic determinants that are responsible for
its phenotype.
Since the bulk amount of cytoplasm containing many
plastids is contributed by the egg and the male gametes
contribute negligible amount of cytoplasm, therefore
plastids present in the cytoplasm of egg is responsible for
the appearance of maternal colour in the offspring and the
failure of male plant to transmit its colour to offspring is
reasonable.
In the offspring from variegated female parents, green,
white and variegated progeny are recovered in variable
proportions. The variegated parent produces three kinds of
egg- some with colourless plastids, some contains only
green plastids, and some are with both chloroplasts and
leucoplasts.
As a result, zygotes derived from these three types of egg
cells will develop into green, white and variegated
offspring’s, respectively.
Inheritance of Lojap Trait in Maize:
In maize plant, iojap is a trait which produces green and
white stripped leaves. This trait is controlled by a recessive
chromosomal gene (ij) when present in homozygous state.
The name iojap was derived from ‘Iowa’ state (USA), the
source of maize strain and japonica, the name of a stripped
variety.
When a normal plant with green leaves used as a female
parent is crossed (Fig. 22.4) with iojap parent, the offspring
will be green leaved:

Again, when a reciprocal cross is made between a normal

green plant (used as male) and iojap plant (used as female).
The offspring will be of three different types:

In iojap plants, green and white stripped trait of leaf is

inherited from the female parent due to maternal
inheritance. It seems that iojap plants contain two types of
plastids— normal green, and abnormal iojap plastids.
During the formation of egg cells plastids are randomly
distributed in the egg cells. If the egg cell receives normal
green plastids it will produce green leaved plants
irrespective of which plant acted as pollen parent. If the egg
cell receives abnormal colourless plastids, it will give rise to
white leaved plants. If the egg cell receives both green and
abnormal plastids it will give rise to plants with green and
white stripped leaves.
If stripped leaved Fx iojap (Ij,ij) as female parent is
crossed with normal green leaved (Ij Ij) as male
parent the following types of offspring are
obtained:
This backcross experiment shows that green males have no
effect upon progeny. The appearance of iojap trait has been
explained by two hypotheses. One hypothesis states that
frequent mutation in the chloroplast genome produces the
abnormal plastids.
Another hypothesis suggests that certain cytoplasmic
elements other than chloroplast mutation bring about the
bleaching of chloroplasts. It is also suggested that a nuclear
gene controls the development of abnormal plastids in the
cytoplasm. So this type of inheritance is a case of inaction
between nuclear and cytoplasmic inheritance.
Extra-Nuclear Inheritance by Mitochondria of
Yeast:
Yeast, Saccharomyces cerevisiae, are unicellular
ascomycetes fungi. In this fungi, sexual reproduction takes
place by the fusion of two somatic cells to form a diploid
zygote nucleus. Next follows two successive nuclear
divisions forming four haploid daughter nuclei, all of which
take part in ascospore formation. Now the mother cell, i.e.,
zygote cell, is called ascus.
The diploid zygote can also be grown vegetatively as a
diploid strain that will later sporulate. Respiration of yeast
cell takes place both aerobically and anaerobically
(fermentation). Certain mutant yeast cells are unable to
utilise oxygen and are comparatively small- sized and slow
growing producing small colonies on agar medium. These
small colonies forming mutant strains of yeast are known as
petites.
In petite strains, the necessary components (cytochrome b,
c1) and some enzymes (cytochrome oxidase a, a3) for aerobic
terminal respiration activity are absent. But these
components are present in the cell of normal strain where
they are associated with the inner membrane of
mitochondria.
Petite strain can be maintained indefinitely in the vegetative
state and can be mated with normal yeast cells. When such
mating are carried out, three petite varieties can be
classified:
Nuclear (segregational) Petites:
When a normal haploid strain of yeast is crossed with a
haploid petite strain, a normal diploid zygote is produced.
The haploid ascospores produced from zygote by
sporulation are segregated in the ratio 1 : 1 (petite : normal).
Hence the result of such cross follows ordinary nuclear
mendelian inheritance (Fig. 22.5).

Neutral Petites:
In this type, only normal wild ascospores are produced from
mattings between petite and normal strain of yeast. The
petite characteristics is absent in the product of segregation.
So it shows the non-Mendelian inheritance. This non-
Mendelian behaviour is very difficult to explain on the basis
of nuclear genes and indicates that such petite
characteristics are caused by extra-nuclear inheritance.
Suppressive Petites:
In this type, all ascospores produced from mating between
normal and petite strain are petite type. Such petites seem
to suppress normal respiratory behaviour and the
suppressive petite factor acts as a dominant. Fig. 22.5 shows
diagrammatic scheme for explaining some differences
between neutral and suppressive petite in terms of DNA.
Therefore, there are two different genetic causes for
respiratory deficiency in yeast. One is nuclear and the other
is extra nuclear. On this basis a neutral petite having the
nuclear gene for normally functioning mitochondria is
crossed with a segregational petite (Fig. 22,6). The diploid
zygote produced from such cross can use the normal
nuclear genes from neutral petite and respire normally.
When such diploid zygotes are grown vegetatively as a
diploid strain, they produce diploid colonies that are of
normal size and respire normally. But when such diploid
zygotes are allowed to sporulate, they undergo meiosis and
produce four haploid ascospores of which two are petites
and other two are normal.
It indicates that normal and petite characteristics segregate
in the 1 : 1 ratios expected from mendelian segregation.
It is noted that the neutral petite contains the normal
nuclear gene for the respiratory enzymes but the
segregational petite does not contain respiratory enzymes,
so it is obvious that the cytoplasmic factor of the neutral
petite appears in the cytoplasm of diploid zygotes where the
factor is possibly independent of nuclear control.
It is also noted that neutral petite strains are readily
produced by subjecting normal strain in low doses of
acriflavines dyes as well as ethidium bromide. But the
treatment of such doses of dye does not induce any nuclear
changes. Thus it strongly indicates the involvement of
extra-nuclear change of gene controlling petite
characteristics. Such changes ultimately shows the extra-
nuclear inheritance of petite characteristics.
(viii) Extra-Nuclear Inheritance by Mitochondria
of Porky Strain of Neurospora:
There are many examples for mitochondrial enzyme
deficiency which are cases of extra chromosomal
inheritance—petite yeast came from the studies of
Neurospora crassa, a member of ascomycetes group of
fungi. In this fungus, there is a slow-growing mutant strain
called porky.
Such strain exhibits poorly differentiated mitochondria
which are deficient in the membrane-bound cytochromes b,
a1 and a3, essential proteins of the respiratory electron
transport chain and also possess greatly reduced numbers
of the small ribosomal units. As in yeast, this trait—
inherited via the female parent In some strains in non-
Mendelian fashion, indicate its extra chromosomal nature.
When porky as female parent (proto perithecial parent) is
crossed with a normal strain as a male parent
(mitochondrial parent), the progeny axe found to be porky.
In reciprocal cross, the progeny are normal. This non-
Mendelian uniparental inheritance suggests that the
cytoplasm of female parent is important because the only
difference between reciprocal crosses is the contribution of
cytoplasm.
Thus nuclear genotype has no effect on this particular
phenotype.
(ix) Extra-Nuclear Inheritance by Symbionts:
There are many cases of cytoplasmic inheritance which are
actually due to the presence of certain intra-cellular
parasites like bacteria, virus particles etc. that make a
symbiotic relationship with the host cell. They are able to
reproduce within the host cell and look like the cytoplasmic
inclusions.
These cytoplasmic symbionts provide some evidences
regarding the cytoplasmic inheritance of the host cell. These
symbionts are variously designated by Greek alphabets as a
(sigma), k (kappa), etc. The various types of extra-nuclear
inheritances due to parasites or symbionts are discussed
next.
Kappa Particles in Paramoecium:
One of the most stricking and spectacular example of
cytoplasmic inheritance due to symbiont bacteria is noted
in the most common ciblate protozoam Paramoecium
aurelia. In 1943, T. M. Sonneborn reported that some
strains of P. aurelia contain kappa particles and are known
as killer strain.
Kappa particles are the symbiont bacteria called
Caedobacter taeniospiralis. The diameter of kappa particles
are about 0.2µ. They are bounded by a membrane and
contain a little bit of cytoplasm with DNA. The strain of
Parameocium in which the kappa particles are absent are
called sensitive strain. The sensitive strains are killed by the
killer strain.
The destruction of sensitive strain occurs through secretion
of a toxic substance called paramecin. This toxic substance
is believed to breakdown the food vacuole membrane of the
sensitive strain. Paramecin is diffusible in the liquid
medium (Fig. 22.8).
When killers are allowed to remain in a medium for a time,
they are not killed. It means that paramecin has no effect on
killers. Paramecin is associated with a particular kind of
kappa that occurs in about 20 percent of a kappa
population.
These kappa bacteria possess a refractile protein containing
‘R’ body and are called brights because they are infected
with a virus that controls the synthesis of a viral protein as
well as R protein body in kappa bacterium.
The virus may act as the toxin in the killing response and R
body facilitates the penetration of the toxin. The non-bright
kappa bacteria may also contain virus but the virus may be
in provirus state in them.
The killer character of Paramoecium has a nuclear as well as
cytoplasmic basis. The existence of kappa particles is
determined by presence of a nuclear dominant gene K.
Kappa particles, like other bacteria, multiply through
fission.
But their multiplication in the cytoplasm of Paramoecium
depends on the presence of a dominant nuclear gene K
which helps to make an environment necessary for the
bacteria to reproduce.
When killer strain of Paramoecium conjugates with
sensitive strain under appropriate condition for brief period
and no cytoplasm exchange occurs, two kinds of clones
result- one from the original killer cell which contains allel
K (Kk) and kappa particles and the other from the original
sensitive cell which carries the allel K (Kk) and lacks kappa
particles.
It indicates that homozygous (either KK or kk) strains
become heterozygous following an exchange of K and k
genes without cytoplasmic exchange.
Following autogamy (a process of self- fertilisation within
one undivided cell resulting in homozygosity), half the
progency (50%) are sensitive Paramecia. But all progenies
of sensitives following autogamy will be sensitive’s.
In this conjugation, following autogamy of killers, 50%
progeny will receive Kk genotype with cytoplasmic kappa
particles other 50% progeny will receive kk genotype with
cytoplasmic kappa particles. But it will be sensitive, because
kappa cannot reproduce in the cells unless a K allele is
present in the nucleus and, as a consequence the kappa are
eliminated.
On the other hand, in this conjugation the product of
autogamy of sensitive strain obtained after conjugation are
all sensitive. All through, 50% progeny of autogamy have
KK genotype without cytoplasmic kappa particles because
no cytoplasm has been transferred in this conjugation.
Remaining 50% progeny of autogamy of sensitive’s have kk
genotype and no cytoplasmic kappa particles.
Under some conditions of conjugation persists much
longer; a long connection is established between conjugants
(killer and sensitive). In this conjugation, cytoplasm as well
as nuclear genes are exchanged (Fig. 22.9). As a
consequence both ex-conjugants will receive the genotype
Kk and the cytoplasm with kappa particles.
Therefore, conjugation for longer period with cytoplasmic
exchange will produce all killer strains. Autogamy of both
ex-conjugants produces homozygotes KK (killer) and kk
(sensitive) cell in the 1 : 1 ratios, respectively, as expected
from Mendelian segregation.
Therefore, conjugation for shorter period without
cytoplasmic exchange does not follow the Mendelian
pattern of inheritance. Hence it confirms the cytoplasmic
basis of inheritance of killer trait.
Mu Particles in Paramoecium:
There is another type of killer trait found in certain strain of
Paramoecium due to presence of ‘mu’ particles in the
cytoplasm. A Paramoecium with a ‘mu’ particle is called
mate killer. On the other hand, a Paramoecium having no
‘mu’ particles is called mate sensitive.
It is so named because when a Paramoecium with ‘mu’ par-
ticle conjugates with a partner Paramoecium without ‘mu’
particle then the former kills the latter. The ‘mu’ particles
exist only in those cells whose micronucleus contains at
least one dominant of either of the two pairs of unlinked
chromosomal genes such as M1 and M2. The ‘mu’ particles
are symbionts which are made of DNA, RNA and other
substances.
The maintenance of the ‘mu’ symbiont in a Paramoecium is
dependent upon the genotype of the Paramoecium. In fact,
the mate-killers of few genotypes maintain their normal
number of particles for about seven generations. From the
eighth generation, the particles suddenly and completely
disappear from the a small fraction of the cell.
Gibson and Beale (1962) suggested that the maintenance of
‘mu’ particle in Paramoecium was due to the presence of
another cytoplasmic particle called metagon. It is possibly a
long-lived messenger RNA or informosome and may be a
product of M1 and M2gene. One metagon may be necessary
for the maintenance of hundred ‘mu’ particles.
(x) Sigma Virus in Drosophila:
Some strains of Drosophila melanogaster are sensitive to
C02 as they die when briefly exposed to C02, while normal
flies can be exposed for long periods to pure C02 without
permanent damage. The high degree of C02– sensitivity is
associated with the presence of a DNA virus called sigma
factor found in the cytoplasm of C02 sensitive Drosophila.
Sigma factor is transmitted through the egg cytoplasm.
When a cross is made between C02– sensitive female with
normal male, all offspring’s are C02 sensitive. Again, in
reciprocal cross, i.e., a cross between normal female and
C02 sensitive male, most of the offspring’s are normal except
for a small proportion of progeny which are C02sensitive.
Therefore, the inheritance pattern of C02 sensitivity is non-
Mendelian and confirms the cytoplasmic basis of inheri-
tance.
(xi) Spirochaetes and Maternal Sex Ratios in
Drosophila:
Spirochaetes sometimes enter into the female body cell of
Drosophila and live there as endoparasites. When
spirochaetes enter the egg cell and these infected egg cells
are fertilised, the zygotes having XY sex. chromosome are
killed early in embryonic development and XX zygotes
survive.
Therefore, the presence of spirochaete in the female body
gives rise to exclusively female progeny—this condition is
known as maternal sex ratio. It is evident that XY embryos
are killed by a toxic substance which may derive from a
DNA virus present within spirochetes that live as
endoparasite in the female body of Drosophila. Maternal
sex ratios in Drosophila is also considered as an example of
extra nuclear inheritance.
Milk Factor in Mice:
This is an interesting example of extra nuclear inheritance.
It is found that certain types of mice are very susceptible to
mammary cancer and this characteristic is found to be
transmitted maternally.The results of reciprocal cross
between susceptible mice and low-incidence mice depend
on the trait of female parent.
When the young mice of low cancer incidence parent are
allowed to feed milk by a susceptible foster mother, it
produce, a high rate of cancer in them. Hence this is a case
of infective agent transmitted in the milk. The milk factor
responsible for causing cancer is possibly a virus. The
presence of milk factor depends on nuclear gene.
(xii) RNA Viruses in Fungi:
Like Paramoecium, there are two strains of yeast
(Saccharomyces cerevisial). One strain is killer and other
one is sensitive. The hiller strain secretes a proteinaceous
toxic substance that kills the sensitive strain of yeast cell.
When a cross is made between killer and sensitive strain of
yeast, only killer offspring’s are produced—indicating
uniparental inheritance. There are some other strains of
yeast which are called neutral strains.
Neutral strains are neither killed by killer nor do they kill
the sensitive strain. But the cytoplasm of both killer and
neutral strains contain two types of double-stranded RNA
in the form of isometric virus-like particles (about 39 nm in
diameter).
The existence and maintenance of virus particles in the
yeast cytoplasm are controlled by some dominant nuclear
genes called MAK genes (maintenance of killer). Some other
nuclear genes—e.g., KEXx (killer expression) and KEX2—
convert killers into neutrals.
A similar situation is noted is case of Ustilago maydis, a
maize smut fungus. Here the cytoplasm of killer strain also
contains maycovirus like particle containing double-
stranded RNA. Killer strain secretes a toxin which kills
sensitive strains but it has no lethal effect on resistant
strains. Resistant strains are particularly resistant to one of
the killer strains designated as p1, p4 and p6. Some nuclear
genes denoted as Pr1,Pr4 and pr6 convert sensitive strain into
resistant ones.
In all such cases mentioned above, the virus like particles
are not the integral part of the normal cellular organisation
but their existence and transmission indirectly provides
some evidences in favour of cytoplasmic inheritance.
(xiii) Uniparental Inheritance in Chlamydomonas
Reinhardi:
R. Sager (1970) and N. Gilham (1968) have reported some
cases of extra-chromosomal inheritance in green alga
Chlamydomonas reinhardi. The alga reproduces by asexual
as well as sexual means. The sexual reproduction takes
place by fusion between two morphologically similar but
physiologically dissimilar haploid gametes coming from
different haploid parents designated as ‘+’ and ‘-‘.
The gametic fusion produces the zygote. The sex is
determined by a single chromosomal gene. When meiosis
occurs in the zygote, four haploid daughter protoplasts are
formed which give rise to a new plant body. Although both
the sexes contribute equally to the zygote, there is maternal
transmission of certain cytoplasmic traits.
Chlamydomonas is a haploid unicellulate green alga. It has
two mating types—’+’ and ‘-‘.The two mating types are
governed by two alleles of a nuclear single gene. The alleles
axe named as mt+ and mt–. The + mating type is considered
as female, while the – mating type is regarded as male.
During sexual reproduction one mt+ and one mt–cell pair
and fuse together to form a zygote where there is mixture of
cytoplasm coming from both mt+ and mt– gametes.
The zygote undergoes meiosis to produce 4 haploid
meiozoospores of which two zoospores contain ‘+’ alleles
and other two contain ‘-‘ alleles, i.e., it shows typical 1 : 1
segregation for nuclear genes. But for their plasma genes all
zoospores are identical and contain only mt+type plasma
genes by mt+ plasma genes. The inactivation is not clear but
it may involve an enzymatic process.
R. Sager isolated two strains of Chlamydomonas: one strain
was resistant (Sr) to 500/xg of streptomycin per ml of
culture solution and the other one is sensitive. The trait of
streptomycin resistance is believed to be located in its cp-
DNA (chloroplast DNA).
Mating between mt+ streptomycin resistant (Sr) and mt
sensitive (Ss) cells produce only resistant progeny but the
nuclear genes for mating type segregate as expected (Fig.
22.10). But the reciprocal cross between mt+ susceptible
and mt– resistant shows again the expected segregation for
mating type but all progenies are sensitive type. Therefore,
it clearly provides an example for extra-nuclear inheritance.
It is also observed that in less than 0.1% of zygotes plasma
genes from mt” parent are not inactivated and produce
cytohets, i.e., heterozygotes for cytoplasmic genes.
(xiv) Inheritance of Preformed Structures:
In some lower group of animals like ciliated protozoa there
are some characteristics which show cytoplasmic
inheritance. These traits are not controlled by any plasma
gene. In Paramoecium, the cytoplasm is differentiated into
cortex or ectoplasm and medulla or endoplasm.
The mouth and the contractile vacuole are the prominent
preformed traits that are present in the cortical region of
Paramoecium. These preformed structures can be transmit-
ted independent of the transmission of nuclear genes and
plasma genes.
For normal sexual reproduction two individuals of
Paramoecium called conjugants come close together,
exchange their nuclear material, and then separate as ex-
conjugants with zygote nucleus. In some rare cases,
conjugants do not separate and remain as doublet animal
with two sets of mouth and contractile vacuole structures.
When this doublet animal reproduces asexually by binary
fission, they also give rise to doublet ex-conjugants. When
doublets are mated with normal singlet’s, the progeny of
doublets ex-conjugants are doublets and the progeny of
singlet’s are singlet’s. This type of inheritance is also found
when the animals reproduces through autogamy.
In these crosses it is noted that the nuclear genes of
doublets and singlet’s are inherited in normal Mendelian
fashion. But the mode of duplicated mouth and contractile
vacuole is independent of the mode of inheritance of
nuclear genes as well as cytoplasmic hereditary factors.
Furthermore, the transplantation experiments reveal that
such cortical structure (mouth, contractile vacuole) are
autonomous and are maintained by cell division.
Some born has suggested that different parts of the cortex
might serve as sites for the specific absorption and
orientation of molecules derived from the milieu and
genetic action. Besides this, preformed cortical structures
could act by determining where , some gene products go in
the cell, how these combine and orient and what they do.
(xv) Extra-Nuclear Inheritance in Bacteria:
In many cases, extra-nuclear inheritance has also been
reported in bacteria. In addition to main chromosome,
bacterial cell has got a unique extra-nuclear genetic system
in their cytoplasm. Such extra-nuclear genetic system plays
an important role for cytoplasmic basis of inheritance.
In bacteria, the cytoplasmic inheritance is
performed by:
(i) plasmids, and
(ii) episome.
In addition to main chromosome, subsidiary DNA is also
present in the bacterial cell in the form of plasmid. Plasmids
are mini-circular DNA duplex and are capable of
independent replication and transmission. By definition, a
plasmid is a relicon that is stably inherited in an extra-
chromosomal state.
Plasmids are not essential for the survival of bacterial cells,
i.e., disposable-except under certain environmental
conditions. Plasmids vary in size and contain three to
several hundred genes. A bacterial cell may contain more
than one plasmid.
There are several types of bacterial plasmids of
which three general types have been studied
extensively such as:
(a) F plasmid;
(b) R plasmids, and
(c) Col plasmids.
F plasmids carry genes for the development of F pili and are
responsible for conjugation.
R plasmids carry genes which are responsible for resistance
to antibiotics or other antibacterial drugs.
Col plasmids carry genes which code for colicins. Colicins
are proteins that kill sensitive E.coli cells.
Plasmids may again be divided into two types on the basis
of whether or not they mediate conjugative self-transfer.
They are:
(a) Conjugative and
(b) Non-conjugative.
Conjugative plasmids are also known as transmissible
plasmids that mediate the transfer of DNA through
conjugation. All F plasmids, R plasmids and some col
plasmids are the examples of conjugative plasmids.
These plasmids spread rapidly among the bacterial cells of a
population. Transmission of R plasmids is responsible for
many pathogenic bacteria to become resistant to many of
the widely used antibiotics such as penicillin, tetracycline,
streptomycin, kanamycin, chloramphenicol etc.
The transmission of these plasmids are not only restricted
among the population of the same species but are also
known to transfer to others like Proteus, Salmonella,
Hemophilus, Pasteurella, Shigella etc.
Non-conjugative or non-transmissible plasmids do not
mediate DNA transfer through conjugation. Some R and Col
plasmids are of this type.
Some plasmids are capable of becoming attached and
integrated into the bacterial chromosome. Then they are
named episomes.
Plasmid and episomes contain insertion sequences which
are also present in bacterial chromosomes. Insertion
sequences are transposable in that they can move about
within and between chromosomes and mediate genetic
recombination between otherwise non-homologous genetic
elements within which they are located.
Insertion sequence are also responsible for the transfer of
genetic controlling resistance to antibiotics from one
genetic element to the other.
Considering the mode of transmission, location and the
presence of genes controlling certain characteristics, it is
clear that the phenomenon of extra-nuclear inheritance still
exists even in most simple, prokaryotic unicellular organism
like bacteria.
Considering the discussion of this article we can
summarise the characteristic features of
cytoplasmic inheritance as:
i. In case of cytoplasmic inheritance, reciprocal crosses
show marked difference for characteristics governed by
plasma genes.
ii. In most cases female parent contributes the plasma
genes, i.e., uniparental inheritance or maternal inheritance.
iii. In general, F2, F3 and so on generations do not show
segregation for cytoplasmically inherited characteristics. It
is a non-Mendelian inheritance.
iv. In case of bi-parental inheritance, irregular segregation
takes place in F1.
v. Several plasma genes are associated with cp-DNA or mt
DNA in higher eukaryotic organisms and with plasmids or
eRisomes in prokaryotes.
vi. In many cases, a cytoplasmically inherited characteristic
is associated with an endosymbiont or parasite or virus
present in the cytoplasm of the organism.
In some specific cases the cytoplasmic inheritance of some
preformed characteristics is not affected by exchange of
cytoplasm and is not controlled by nuclear genes. They are
autonomous and are maintained by cell division.