Extra-chromosomal inheritance

Contents: Definition; Chromosomal (nuclear) versus cytoplasmic inheritance; Types of

extra-chromosomal inheritance; Extra-chromosomal inheritance in Paramecium: Strains of
Paramecium; Killer trait in Paramecium; Properties of kappa particles; Extra-chromosomal
inheritance in Paramecium; Extra-chromosomal inheritance in Drosophila; Suggested
reading.

Definition
Inheritance of certain traits of organisms that are controlled by the genetic materials present
in the cytoplasm is known as cytoplasmic inheritance, which is also referred to as extra-
chromosomal or non- nuclear inheritance. The genetic materials present in the cytoplasm are
called ‘plasmagenes’.

Chromosomal (nuclear) genes versus plasmagenes

Fig 12.1 Chromosomal (or nuclear) genes= DNA and RNA in the chromosomes;
Plasmagenes= mitochondrial DNA (mtDNA) and chloroplast DNA (cpDNA) in the
cytoplasm

Differences between chromosomal (nuclear) and cytoplasmic inheritance

Chromosomal inheritance Cytoplasmic inheritance

1. Traits are controlled by nuclear genes Traits are controlled by plasmagenes present
present in the nucleus. in the cytoplasm.
2. Except for sex-linked traits, father and The role of mother is more important than that
mother play equal role. of father, because ova (eggs) contain more
cytoplasm than that of spermatozoa (sperms).
3. Reciprocal crosses between males and Reciprocal crosses between males and
females produce Mendelian ratios. females produce non-Mendelian ratios.
4. Traits are transmitted through Traits are transmitted through cytoplasm.
chromosomes.

Lecture 12_Extra-chromosomal inheritance 1

Types of extra-chromosomal inheritance
There are three major types of extra-chromosomal inheritance as follows:
1. Infectious particles: Killer trait in Paramecium aurelia, CO2 sensitivity in
Drosophila melanogaster, milk factor in Mus musculus etc.
2. Cell organelles: Plastid (chloroplast) transmission in 4 o’clock plant, Mirabilis
jalapa.
3. Maternal factors: Shell coiling in land snail, Limnaea peregra, eye colour variation
in flour moth, Ephestia cautella etc.

Infectious particles other than kappa (ϰ):

(a) μ (mu) particles: Produce mate killer and mate sensitive strains;
(b) π (pi) particles: Mutant forms of kappa and do not produce poisonous substance;
(c) λ (lambda) particles: Cause lysis or disintegration of sensitive strains;
(d) σ (Sigma) particles: Cause unconsciousness to the sensitive strains;
(e) DNA viruses: Cause breast cancer (mammary carcinomata).

Extra-chromosomal inheritance in Paramecium

Extra-chromosomal or cytoplasmic inheritance in Paramecium was studied extensively by
the American scientist T. M. Sonneborn and his colleagues at the University of Indiana, USA.
They studied the strains of Paramecium, their properties and the mode of transmission of the
extra-chromosomal substance the then called kappa particles in this ciliate protozoan.

Fig 12.2 T. M. Sonneborn (1965) who contributed to our understanding of the extra-
chromosomal inheritance in Paramecium
Strains of Paramecium
There are two strains of Paramecium, characteristics of which are as follows:

1. The killer strain: Cytoplasm contains toxic and infectious particles called kappa, which
produces a water soluble chemical (paramecin); cytoplasm is opaque and particulate; bears
nuclear dominant genotypes KK (homozygous) or Kk (heterozygous).

2. The sensitive strain: Absence of kappa particles, so, cytoplasm does not contain any toxic
or infectious particles and hence unable to produce paramecin; cytoplasm is transparent and
non-particulate; bears nuclear recessive genotype kk.

Lecture 12_Extra-chromosomal inheritance 2

 Fig 12.3 The killer (left) and sensitive strains of Paramecium
Killer trait in Paramecium
 Certain Paramecium secretes toxic substance (paramecin) into the medium around
them, which kill other paramecia;
 This killer trait is caused by kappa particles, which are symbiotic bacteria
Caedobacter taeniospiralis (meaning killer bacterium with spiral ribbon);
 The kappa particles are found in natural populations of Paramecium aurelia.

Properties of kappa particles

1. Each kappa particle is about 0.2μ in size.
2. It undergoes cell division, i.e. it can multiply.
3. Killer strains may harbour up to 1600 kappa particles.
4. To be a killer, at least 400 kappa particles need to be present in the cytoplasm.
5. EM studies revealed that kappa particles are actually symbiotic bacterium,
Caedobacter taeniospiralis.
6. Kappa particles can be destroyed by the physical and chemical mutagens like
low temperature, X-rays, nitrogen mustard etc.

Extra-chromosomal inheritance in Paramecium

Two types of reproduction take place in Paramecium (Fig. 12.4). These are:
(1) Asexual reproduction by binary fission; and
(2) Sexual reproduction by conjugation.

Conjugation is again of two types:

(1) Normal conjugation: Usual method; exchange of nuclear materials only; No
cytoplasmic exchange takes place; so, exconjugant killer produces 50% killer and
50% sensitive; but exconjugant sensitive produces sensitives only. So,
transmission of kappa particles through the killer strain does not take place.

(2) Rare conjugation: Unusual method; a cytoplasmic bridge is formed; exchange of

both nuclear and cytoplasmic materials takes place; exconjugant killer produces 50%
killer and 50% sensitive, whereas exconjugant sensitive produces 50% sensitive and
50% killer. That means, transmission of kappa particles through the killer strain takes
place only through the rare conjugation.

Lecture 12_Extra-chromosomal inheritance 3

 Fig 12.4 Reproduction in Paramecium by conjugation and binary fission

Normal and rare conjugations in Paramecium

In normal conjugations, when a killer strain conjugates with a sensitive strain, the resulting
exconjugants are similar to their parents, both killer and sensitive, because the conjugation
takes place for a brief period of time and the exchange of cytoplasmic materials does not
occur. Then the exconjugants further divide asexually by binary fission. Here the sensitive
strain produces only sensitives, but the killer strain produces both killer and sensitive strains
because of the further dilution of kappa particles (Fig. 12.5 left).

In rare conjugations, however, when a killer strain conjugates with a sensitive strain, the
resulting exconjugants are both killers, because the conjugation takes place for a long period
of time and the exchange of cytoplasmic materials including kappa particles occurs. Finally,
the exconjugants further divide asexually by binary fission, when the killer strain produces
both killer and sensitive strains (Fig. 12.5 right).

Fig 12.5 Consequences of normal (left) and rare (right) conjugation in transmitting killer trait
(kappa particles) in Paramecium

Lecture 12_Extra-chromosomal inheritance 4

Conclusions
Extra-chromosomal inheritance of the killer trait (kappa particles) in Paramecium is
dependent on the following three important conditions:
1. The Paramecium must carry KK or Kk (i.e. K-) nuclear genotype;
2. At least 400 kappa particles must be present in the cytoplasm; and
3. The kappa particles are transferred from the killer strain to the sensitive strain by rare
conjugation, in which a cytoplasmic bridge is formed; the conjugation takes a longer period
of time; and both nuclear and cytoplasmic materials are transferred.

Extra-chromosomal inheritance in Drosophila

The following example shows the inheritance of extra-chromosomal substance (called sigma
particles) in the fruit fly, Drosophila.

Fig 12.6 The fruit fly, Drosophila melanogaster shows an extra-chromosomal inheritance of
sigma particles

 Most Drosophila can tolerate an exposure of pure CO2 for long hours without any
injury.
 French geneticists Ph. L’Heritier and G. Teissier (1937) discovered a strain of
Drosophila melanogaster which was sensitive to CO2.
 The sensitive flies, when exposed to CO2 for a short period, become unconscious in a
characteristic way, with their legs becoming paralyzed, and eventually they die.
 A sensitive fly retains its sensitivity trait, even when all its chromosomes are replaced
by those of the normal fly, suggesting that the causal agent is located in the
cytoplasm.
 Later on, the causal agent has been called sigma (σ) particles.

Characteristics of sigma (σ) particles

1. Sigma particles are DNA viruses, each particle is of ~0.07μm in diameter and are
responsible for their hereditary transmission;
2. They transmit through egg cytoplasm, but occasionally through sperms;
3. Their reproduction depends on suitable temperature of 20°C;
4. They are heat labile, for example, destroyed by high temperatures;
5. They show properties of non-chromosomal gene or plasma gene.

Lecture 12_Extra-chromosomal inheritance 5

Crossing experiments on CO2 sensitivity in Drosophila
The crossing experiments with Drosophila revealed the following results:
1. P: Normal ♀♀ × Sensitive ♂♂
F1: Most offspring are normal
(Only a few progeny are sensitive)

2. P: Sensitive ♀♀ × Normal ♂♂
F1: All offspring are sensitive

3. P: Cell-free extracts from sensitive ♀♀ → When injected into normal ♀♀

F1: All offspring are sensitive

Conclusions:
(a) The inheritance pattern of CO2 sensitivity in Drosophila is non-Mendelian
(Experiments 1 and 2);
(b) The CO2 sensitivity trait shows cytoplasmic basis of inheritance (Experiment 3).

Suggested reading:
Burns, GW. 1980.
Gardner et al. 1991.
Islam, MS. (2018).
Sinnott et al. 1973.
Winchester, AM. 1966.
Wikipedia: www.wikipedia.com.
Bmjvg, g.mv. I Ab¨vb¨ 2017|

Lecture 12_Extra-chromosomal inheritance 6