Extrachromosomal Inheritance-

Definition, Criteria, Maternal

Inheritance and Examples
“A non-mendelian pattern of inheritance governed by the DNA present in the

cytoplasm is known as extrachromosomal inheritance or cytoplasmic inheritance.”

The DNA is the genetic material of us and arranged on chromosomes. It helps to store

and transfer information (called traits) through the process of replication, transcription

and translation. Nuclear DNA is the basis for inheritance of almost all type of

phenotype of ours. It inherited in a particular fashion from parents to their offspring.

Though all genes are inherited in Mendelian style, some genes present in the

cytoplasm of the cell, inherited in a non-mendelian pattern. This type of inheritance is

called as extrachromosomal inheritance or cytoplasmic inheritance.In the present

article; we will discuss one of the amazing topic- “extrachromosomal inheritance.”

What is extrachromosomal inheritance?

The extrachromosomal inheritance also is known as cytoplasmic inheritance or non-

mendelian inheritance was first reported by Boris Ephrussi in yeast during 1949.

Cytoplasmic DNA or extrachromosomal DNA is present significantly in some

important organelles like chloroplast and mitochondria. It is a big mystery that how

actually these organelles created their own genome. One theory which stated that it

was a symbiotic relationship. It is believed that mitochondria were once free-living

bacteria. Over a period of time, it created a symbiotic relationship with eukaryotic

cells and established them into the cytoplasm and ultimately evolved as an organelle

in living eukaryotic cell.

Similarly, the chloroplast in green plants comes from the free-living algae and

established a symbiotic relationship with eukaryotic plant cells and settled into

cytoplast of green plants. Both types of sub-genome have well-developed DNA

machinery which is equipped with the entire component required for central dogma.

Additionally, chloroplast has antibiotic resistance genes indicate that it was derived

from bacteria, previously.

The genome is made up of few genes and several thousand base pairs, still it has their

own rRNA, tRNA and DNA for replication, translation and transcription.

Definition:

“The extrachromosomal DNA present in the cytoplasm and not on chromosomes

which follows the non-Mendelian pattern of inheritance is known as

extrachromosomal inheritance.”

Criteria for extrachromosomal inheritance:

The extrachromosomal DNA follows a non-mendelian pattern of inheritance unlike

the common Mendelian segregation pattern is not observed in the extrachromosomal

DNA because it does not have the centromere it can not segregate, unlike the normal

nuclear DNA.
Their own machinery for protein synthesis:

Unlike nuclear DNA, the organelle DNA or the extrachromosomal DNA has its own

replication and transcription machinery. It synthesised their won DNA.

Maternal inheritance:

The extrachromosomal DNA inherited from the maternal side.

The segregation is observed in somatic cells rather than germ cells, unlike

nuclear inheritance.

Examples:
Carl Correns in 1908, first reported non-mendelian inheritance in Mirabilis

Jalapa plastid DNA. Another extrachromosomal inheritance was reported by M M.

Rhoades in 1933. He postulated that inheritance of male sterility in maize is governed

by maternal inheritance and it becomes one of the greatest discoveries in science.

Another important point that makes extrachromosomal DNA even unique is maternal

inheritance. It inherits from mother to their offspring which means that only female

individual from the entire population can inherit cytoplasmic DNA.

One theory suggests that female reproductive cell (ovum) is bigger, contain more

cytoplasm and more organelles than male reproductive cells. This would be expected

to influence non-mendelian inheritance or maternal inheritance.

The maternal-effect in snail:
The character of coiling in snail is governed by maternal inheritance.
Snail, Limnaea peregra, has two types of shell coiling phenotypes: one is dextral
shells which coil for the right side and another is a sinistral shell which coils for
the left side.
Here, the mother’s genotype (not a phenotype) is exclusively responsible for the
development of coiling style. Assume that D+ genotype codes for dextral (right
side) coiling and D is codes for sinistral coiling. The reciprocal cross of D+ and D
is shown in the figure:

The image represents the maternal effect of the snail. Here crossing between dextral female and sinistral male results in
dextral offspring in F1, while inbreeding results in all dextral progenies in the F2 generation.

Crossing between D+D+ female and DD male, all the F1, as well as F2 progeny,
become dextral as the mother is D+D+ dextral, here the DD recessive phenotype is
not expressed and typical Mendelian 3:1 ratio is not obtained (all four are
dextral).
In another condition when DD sinistral female is crossed with D+D+ dextral male,
F1 offspring become sinistral with genotype D+D, here mentioning genotype is
important because the inheritance is governed by genotype not by phenotype.
When this F1 progeny is inbred (D+D * D+D) all the F2 progeny become dextral
and coil for the right side. This results indicated that phenotype of parents do not
have any influence on the phenotype of progeny because although all of the F1
progeny are sinistral, all F2 offspring becomes dextral.

The image represents the maternal effect in the snail. Crossing between sinistral female and dextral male results in
sinistral F1 progenies. Though all F1 progenies are sinistral, all F2 progenies become dextral.

Detailed investigation shows that spindle formed during the second metaphase
division decides the direction of coiling. The spindles of dextral snail are tipped to
right and vice verse for sinistral. Interestingly, spindle arrangement in metaphase
in controlled by maternal genes.
So the actual phenotype of “type of coiling” in snail is governed by maternal
genes and it does not depend on the phenotype of any of parent.
In some of the organism, the amount of exchange of cytoplasm plays a crucial role
in the inheritance of phenotype.
Inheritance of kappa particles in paramecium:
Paramecin is a substance found in some of the killer strain of paramecium which
kills the sensitive strains. Paramecin production is governed by the kappa particles
present in the cytoplasm of the paramecium.

The image represents inheritance of kappa particles in paramecium at a shorter period of conjugation.

Here KK gene is responsible for the production of kappa particle which is

dominant over kk gene. In case of inheritance of kappa particle, cytoplasmic
exchange during conjugation plays a crucial role.
When KK killer strains are crossed with kk strains by conjugation, all the progeny
obtained are heterozygous with genotype Kk but the phenotype of
paramecium depends on presence or absence of kappa particles and it will be
influenced by time of conjugation.
The image represents inheritance of kappa particles in paramecium at the longer period of conjugation which results in
the exchange of cytoplasm.

If both are conjugates for a shorter period of time, in F1 generation killer strains
remain killer and non-killer remain non-killer in the heterozygous condition. Here
only nuclear genes are transferred but the cytoplasm is not exchanged between
both strains.
In another condition, if killer and non-killer strains are conjugated for a longer
period of time, due to the exchange of kappa particles, sensitive strain receives
kappa particles through cytoplasmic exchange and sensitive strains become killer
in F1 generation.
Variegation in Four O’clock Plant:
The classic study of maternal inheritance was performed by Correns on the four o'clock plant
(Mirabilis jalapa). This plant can have either green, variegated (white and green) or white
leaves. Flower structures can develop at different locations on the plant and the flower color
corresponds to the leaf color. When Correns crossed the different colored flowers from different
locations on the female plant with pollen obtained from flowers of the three different colors, the
progeny that resulted from the cross always exhibited the color of the leaf of the female. That is,
regardless of whether the pollen was from a leaf that was green, variegated or white. If the
female flower came from a region where the leaves where green, all the progeny were green.
Similar results were seen when the female was from a region on the plant where the leaves were
either variegated or white. In comparison to traits controlled by maternal effects, those traits
controlled by maternal inheritance, the female phenotype is always expressed in its offspring.

Female Male Progeny Phenotype

Green Green, Variegated or White Green
Variegated Green, Variegated or White Variegated
White Green, Variegated or White White
Ovules derived from fully green portions of the plant, regardless of the source of pollen, will
result only fully green plants and variegated character will not reappear in the subsequent
generations.

When ovules are derived from variegated branches, three types of seeds are produced in variable
numbers, again regardless of the male parent; some give rise to (sure green, some to pure white,
and the majority to variegated offspring’s. Ovules derived from pale branches, will result only
pale plants.

The results can be explained in the following manner. All of the organelle DNA that is found in
an embryo is from the female. The egg cell is many times larger than the pollen cells, and
contain both mitochondria and chloroplasts. Pollen is small and is essentially devoid of
organelles, and thus organelle DNA. So any trait that is encoded by the organelle DNA will be
contributed by the female. In the case of the four o'clock plant, the different colors of the leaves
is a result of the presence or absence of chlorophyll in the chloroplast, a trait that can be
controlled by the chloroplast DNA. Thus, green shoots contain chloroplasts that have
chlorophyll, the chloroplasts in the white shoots contain no chlorophyll, and the variegated
shoots contain some chloroplasts with chlorophyll and some without chlorophyll. Thus,
depending upon the location in the plant where the flower comes from, the egg can have
chloroplast with chlorophyll, without chlorophyll or a mixture of the two types of chloroplasts.
This is the biological basis of maternal inheritance.