CYTOPLASMIC INHERITANCE

MENDELIAN INHERITANCE:-
Inheritance of most of the characters in eukaryotic organisms shows the following
characteristic features.
1. The contributions by both male and female parents are equal so that the results from
reciprocal crosses are identical.
2. Segregation produces the characteristic 3:1 ratio in the F2 generation of a monohybrid
cross and a typical 9:3:3:1in dihybrid crosses.
These features of inheritance were first demonstrated by Mendel: consequently,
such an inheritance pattern is referred to as Mendelian inheritance. It is universally accepted
that genes showing Mendelian inheritance are located in the chromosomes of eukaryotic
nuclei. Therefore Mendelian inheritance pattern is regarded as a sufficient evidence for a gene
to be located in the chromosomes, such genes are termed as nuclear genes or more commonly
simply as genes.

NON MENDELIAN INHERITANCE

But some characters in several organisms do not show Mendelian inheritance or they show a
non Mendelian inheritance pattern. In such cases, the following characteristic features are
observed.
1. There is consistent difference between the results from reciprocal crosses; generally only
the trait from female parent is transmitted.
2. In most cases, there is no segregation in the F2 and subsequent generations.
Characters showing non Mendelian inheritance may be grouped under 3 broad categories:
(1) those related to cellular structures and patterns,
(2) those produced by intracellular parasites, symbionts and viruses
(3) those associated with DNA containing cell organelles viz., mitochondria and chloroplasts.
Inheritance due to genes located in cytoplasm (plasmagenes) is called Cytoplasmic

inheritance. Since genes governing traits showing Cytoplasmic inheritance are located outside
the nucleus and in the cytoplasm, they are referred to as plasma genes.
The sum total of genes present in the cytoplasm of a cell or an individual is known as plasmon
while all the genes present in a plastid constitute a plastron . The plasma
genes are located in DNA present in mitochondria (mt DNA) and in chloroplasts (cp DNA).
Together both the DNAs are called organelle DNA. Therefore, this type of inheritance is often
referred to as organellar inheritance, plastid inheritance or mitochondrial inheritance.

In this generally, the character of only one of the two parents (usually female) is transmitted to
the progeny. Hence such inheritance is usually referred to as extra-nuclear or extra-
chromosomal or maternal or uniparental inheritance.
The cytoplasmic inheritance is of two types: 1) Plastid inheritance and 2)mitochondrial
inheritance.
1. Plastidial or chloroplast inheritance: Plastids self duplicated and have some amount of
DNA and plays an important role in Cytoplasmic inheritance. Plastids have green pigments
called chloroplasts.
Chloroplasts contain a unique circular DNA (cp DNA) in the stroma that is completely different
from the nuclear genome. Some examples of plastid inheritance are given below:

a) Leaf variegation in Mirabilis jalapa :

 The conclusive evidence for cytoplasmic inheritance was first presented by C. Correns in
Mirabilis jalapa (Four ‘O’ clock plant) in 1909. He studied inheritance of leaf variegation in
M. jalapa. Variegation refers to the presence of white or yellow spots of variable size on the
green background of leaves.
 In M.jalapa, leaves may be green, white or variegated. Some branches may have only green,
only white or only variegated leaves.
 Correns made crosses in all possible combinations among the flowers produced on these 3
types of branches. When flowers from green branch were used as female parent, all the
progeny were green irrespective of the phenotype (green, white or variegated) of male
parent.
 Similarly, progeny from crosses involving flowers bloomed on white branches as female
parent were all white irrespective of the phenotype of male parent. But in progeny from all
crosses involving flowers born on variegated branches as female parent, all the three types
i.e. green, white and variegated individuals were recovered in variable proportions.

Female Parent Male Parent Type of leaf in the progeny grown from
seed

Green Green Green

White
Variegated
White Green white
White
Variegated
Variegated Green Green, white and variegated
White in variable ratio in each of the
Variegated cases.

The green leaf branches have normal chloroplasts, white branches have mutant chloroplasts
and variegated have a mixture of both normal and mutant chloroplasts. The above results
indicated that the inheritance is governed by chloroplasts. Since the cytoplasm is contributed to
the zygote mainly by female parent, the plastids are transmitted to the zygote from the female
parent. Thus the plastids are responsible for variation in the crosses of green, white and
variegated leaves.
b) Maternal inheritance by ‘iojap’ gene in maize
Rhoades (1946) identified the ‘iojap’ gene (ijij) in maize located in chromosome VII controlling plastid
inheritance in the plant. The gene ‘Ij’ is responsible for the normal green colour of the plant.
 In maize, there are three types of leaves i.e. green, iojap (green and white stripes) and white.
The green leaves have normal plastids. Iojap leaves have a mixture of normal and mutant
plastids and white leaves have only mutant plastids. In a cross between green female and iojap
male, only green individuals are produced in F1 generation. But in the reciprocal cross (iojap
female and green male) all the three kinds of progeny are obtained in variable proportions in F1.
Parents Female x Male
Phenotype Green x Iojap
Ij Ij ↓ ijij
Generation F1 Green (Ijij)

Reciprocal cross
Parents Female x Male
Phenotype Iojap x Green
↓

Generation F1
Green, Iojap, White in variable
proportion
These reciprocal differences can be attributed to the type of plastids in the egg cell since only
female parent is contributing cytoplasm to the zygote.

2. Mitochondrial inheritance: The inheritance of some characters, such as

cytoplasmic male sterility in plants, pokyness in Neurospora etc., is governed by
mitochondrial DNA (mtDNA).
a) Cytoplasmic Male Sterility (CMS) in maize : In several crops, cytoplasmic control of male
sterility is known. In maize, cytoplasmic male sterility (CMS) is governed by mitochondrial DNA.
In such cases, if female parent is male sterile, F1 progeny also will be male sterile, because
cytoplasm is mainly derived from female parent.

b)Pokyness in Neurospora : Neurospora, which is a breadmold has two strains i.e. wild and
poky. The wild strain has normal growth. While the poky which is a mutant has very slow
growth.
A cross between a poky female and a wild male produce only poky progeny. In reciprocal cross
(a cross between wild female and poky male) all the progeny would be wild.
This suggests the presence of cytoplasmic inheritance because only difference between the
reciprocal crosses is in the main contributor of cytoplasm.
Reciprocal crosses
Poky x Wild Wild x Poky
↓ ↓
Poky Wild
3) Non Mendelian inheritance due to parasites,symbionts and viruses:-

Inheritance of Kappa particles in Paramecium:-

In Paramecium aurelia, two strains of individuals have been reported. Kappa particles are called
either bright or non bright depending on their appearance under light microscope.
Bright kappa particle is called as ‘ Killer’ which secretes a toxic substance ‘ paramecin’ and the lighter
strain in known as ‘ sensitive’ and is killed if comes in contact with the ‘paramecin’.
Bright kappa contain a DNA virus, which is essential for toxin production.The kappa is a symbiont (a
gram negative bacterium, caedibacter )living in the cytoplasm and not a normal component of
paramecium cells.
In the cytoplasm of the killer strain the kappa particles (cytoplasmic – DNA) are present and are
absent in sensitive strains. The transmission of kappa particles is through cytoplasm but maintenance of
kappa particles and production of paramecin is controlled by ‘k’ allele .
The killer strains carry dominant allele ‘KK or Kk’ and that sensitive ‘kk’.

Kappa particle shows Cytoplasmic transmission like plasma genes. Generally during conjugation
there is exchange of nuclei but no Cytoplasmic exchange.Therefore during conjugation between killer
and sensitive paramecium the sensitive cell do not receive any kappa particles. As a result sensitive cell
remain sensitive after conjugation.

But occasionally Cytoplasmic exchange does occur during conjugation,in such case both the
strain receive kappa particle. But subsequent propagation of kappa depends on the nuclear genotype of
the strains. kappa particle like other bacteria multiply through fission. However kappa particle are able
to multiply only in animal having the genotype KK or Kk. As a consequences they are eliminated from
such animals. The killer animal with kk genotype ultimately become sensitive due to loss of
their kappa particles.

Characteristic features of cytoplasmic inheritance:

1. Reciprocal difference : Reciprocal crosses show marked differences for characters governed
by plasmagenes. In most cases, plasmagenes from only female parent are transmitted and
hence this phenomenon is also called uniparental inheritance.
2. Lack of segregation : In general, F1, F2, F3 and subsequent generations do not show
segregation for a cytoplasmically inherited trait, as F1 individuals receive plasmagenes from
female parent only.
3. Irregular segregation in biparental inheritance: In some cases, plasma genes from both the
parents are transmitted to the progeny, this is known as biparental inheritance
3. Somatic segregation : Plasmagenes generally show the features in somatic tissues such as
leaf variegation features which is of rare occurrence in case of nuclear genes.
4. Association with organelle DNA: Several plasmagenes have been shown to be associated
either with chloroplast or mitochondrial DNA.
For example: Cytoplasmic Male Sterility (CMS) in sorghum and maize is associated with
mitochondrial DNA.
5. Nuclear transplantation: Nuclear transplantation means nucleus of a cell is removed and
replaced by nucleus of another genotype from a different cell. If nuclear transplantation reveals
a trait to be governed by genotype of cytoplasm and not by that of nucleus, it clearly indicates
that the trait or character is governed by cytoplasmic inheritance.
6. Mutagenesis:Some mutagens are highly specific mutagens which act only on the
plasmagenes and do not affect nuclear genes Eg; ethidium bromide

Induction of mutations by such agents or chemicals in a gene clearly indicates that it is a

plasmagene.

7. Lack of chromosomal location: In many organisms extensive linkage maps of nuclear genes
are available. If a gene is shown to be located in one of these linkage groups, obviously it
cannot be a plasmagene.
8. Transfer of nuclear genome through back crosses: Nucleus of a variety or species may be
transferred into cytoplasm of another variety or species through repeated back crossing with
former, which is used as recurrent male parent. Lines produced in this way are called
alloplasmic lines, since they have cytoplasm and nucleus from different species.
9. Lack of association with a parasite or symbiont or virus : In many cases, a cytoplasmically
inherited character is associated with a parasite, symbiont or virus present in the cytoplasm of
the organism. Such cases cannot be regarded as cases of cytoplamic inheritance. Only those
cytoplasmically inherited characters which are not associated with parasites, symbionts or
viruses can be regarded as governed by plasma genes.
Significance of Cytoplasmic Inheritance
1. Development of cytoplasmic male sterility several crop plants like maize. Pearl millet, sorghum,
cotton etc.
2. Role of mitochondria in the manifestation of heterosis.
3. Mutation of chloroplast DNA and mitochondrial DNA leads to generation of new variation
Differences between chromosomal (nuclear) and extra-chromosomal
(cytoplasmic or extra-nuclear or maternal) inheritance

S.No. Character Chromosomal Extra-chromosomal

inheritance inheritance
1 Location of hereditary Nucleus Cytoplasm
factors
2 Associated with Chromosomes Chloroplasts and
mitochondria
3 Pattern of Inheritance Can be explained by Cannot be explained
mendelism by mendelism
4 Individual hereditary Genes Plasmagenes
factors are known as
5 Hereditary factors are Genome Plasmon
collectively known as
6 Characters of F1 May show dominance or Exhibits only the
progeny may be intermediate characteristic of the
between the parents female parent
7 Reciprocal differences Not observed Observed
8 Segregation of factors Present Absent
and recombination
9 Attributes of progeny Under the control of their Under the control of
own genes cytoplasm of female
parent
10 Action of mutagen Non-specific Very specific
11 Frequency of occurrence Most common Rare
12 Gene mapping Easy Difficult