There are many exceptions to the rule in

genetics.

One of them is that not all inherited

characters are determined by genes located
in the nucleus.
A small minority are controlled by genes
located in cell organelles in the cytoplasm
i.e. cytoplasmic genes, and these of
course are exceptions to the chromosome
theory of inheritance.

Since they are

extrachromosomal/extranuclear (i.e.
outside the chromosomes), such genes are
not subject to the normal rules of
Mendelian heredity.
In the recent past, importance on looking
at genes as the unit of selection has
focused almost exclusively on nuclear
genes.

It has been neglected that an important

component of hereditary material in
organisms is non-nuclear.
This non nuclear pattern of inheritance
could be at variance with nuclear pattern of
inheritance.

This makes a potential for conflict between

nuclear and cytoplasmic genes particularly
w.r.t sex, reproduction, the allocation of
parental investments, and altruism towards
kin.
It is regular part of the life of every
eukaryotic organism(as well as a large
proportion of prokaryotes).
The visibility of chromosomes became an
enormous support in the investigation of
nuclear genetics.

However a relative undetectability of many

cytoplasmic factors caused a severe
handicap in the development of study of
extranuclear inheritance.
But recent proliferation of sophisticated
laboratory techniques has removed these
difficulties and slow developmental study
of cytoplasmic genes has been
complemented with an enormous amount
of research data.
By the mid sixties, there were several
hundreds well authenticated cases of
cytoplasmic inheritance.

But now they number in thousands.

By the mid sixties, there were several
hundreds well authenticated cases of
cytoplasmic inheritance.

But now they number in thousands.

After rediscovering Mendel's laws of
heredity(nuclear/chromosomal
inheritance),

Carl Correns conducted experiments with

the four O’ clock (Mirabilis jalapa) to
explore apparent counterexamples to
Mendel's laws in the heredity of variegated
(green and white mottled) leaf color.
Variegated plants have some branches
which carry normal green leaves, some
branches with variegated leaves (mosaic of
green and white patches) and some
branches which have all white leaves
Seed produced by flowers carried on the
green branches gave progeny which were all
normal green.

Regardless of the fact, whether the

phenotype of the branch carried the flower
used for pollen was green, white or
variegated
Seed taken from white branches gave all
white progeny, regardless of the pollen
donor phenotype.

Seeds from flowers on variegated branches

gave three kinds of progeny, green, white
and variegated, in varying proportions;
again regardless of the pollen donor
phenotype.
In other words, the phenotype of the
progeny always resembled the female
parent and the male made no contribution
at all to the character.

The effect is seen quite clearly in the

difference found between reciprocal
crosses
The genes concerned are located in the
chloroplasts within the cytoplasm, not in
the nucleus, and are therefore transmitted
only through the female parent.
In eukaryote organisms, the zygote
normally receives the bulk of its cytoplasm
from the egg cell and the male gamete
contributes little more than a nucleus.

Any genes contained in the cell organelles

of the cytoplasm will therefore show
maternal inheritance.
In eukaryote organisms, the zygote
normally receives the bulk of its cytoplasm
from the egg cell and the male gamete
contributes little more than a nucleus.

Any genes contained in the cell organelles

of the cytoplasm will therefore show
maternal inheritance.
The leaf variegation is due to two kinds of
chloroplasts: normal green ones and
defective ones lacking in chlorophyll
pigment.

Chloroplasts are genetically autonomous

(i.e. self-determining) and have their own
system of heredity in the form of
chloroplast ‘chromosomes’.
Chloroplast chromosome are small circular
naked DNA molecules which carry genes
controlling some aspects of chloroplast
structure and function.

A mutation in one of these genes affects the

synthesis of chlorophyll, therefore follow
the chloroplast in its transmission and will
not be inherited in the same way as a
nuclear gene.
Another distinction of the inheritance of
chloroplasts is that they have no regular
means of distribution,

such as chromosomes do at mitosis, where

they can be equally shared out to the
daughter cells following division.
A plant that begins life as a zygote
containing a mixture of normal and
mutant chloroplasts cannot therefore
maintain the same mixture in all of its
somatic cells.

Some branches of variegated plants may

therefore remain mosaic while others, by
chance, may turn out to contain all white or
all green chloroplasts in all of their cells.
The chloroplasts of plants carry their
genetic information in the form of small
circular DNA molecules, similar in size and
form to the chromosomes of bacteria.
These DNA molecules contain genes which
code for some of the proteins and RNAs
used in chloroplast structure and function;

It is mutations in these genes which are

most likely to be responsible for the leaf
variegation effects
Chloroplasts are not totally independent of
the nucleus in their heredity

Most of their proteins are coded by nuclear

genes, and mutations in these show normal
Mendelian patterns of inheritance.
The really surprising thing about the
chloroplast DNA is the large number of
copies which are present: up to 300 in a
mature plastid.

Since an average of 160 chloroplasts are

present in a mesophyll cell of the mature
leaf of a cereal such as wheat, this means
that there may be as many as 48 000
chloroplast ‘chromosomes’ per mesophyll
cell.
The chloroplast chromosome has variable
length of DNA base pairs.

The chloroplast ‘chromosome’ has a length

of 155 000 base pairs of DNA in lettuce.
Leaf variegation due to chloroplast
mutation is known in numerous other
genera of plants:

Epilobium and Pelargonium

Many of the other examples of cytoplasmic
inheritance, in a variety of species, appear
to involve characters which are associated
with functions of the mitochondria.

They have to deal with defects in growth

and ATP energy metabolism.

Examples are slow growing mutants in the

fungus Neurospora and mutants in yeast.
The mitochondria, like the chloroplasts,
are self-replicating organelles which
contain their own genes and have a limited
number of characters which are
independent of the nucleus.

They are transmitted mainly through the

female line and mutations in their genes
show the same pattern or maternal
inheritance.
Mitochondrial ‘chromosomes’ have a
similar circular configuration of ‘naked’
DNA as chloroplasts.

In a typical haploid yeast cell each of the

mitochondria contains in the region of 50
small circular ‘chromosomes’.