EXTRA NUCLEAR

INHERITANCE
 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 plasmagenes.

 The plasmagenes 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 are 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.
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 three 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.
 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.
 Cytoplasmic Inheritance Involving Dispensable Hereditary
Units (nuclear genes may or may not be involved)
Kappa particles in Paramecium
 Kappa particles are found in certain killer strains of Paramecium and are responsible
for production of substance paramecin, which is toxic to strains not
possessing kappa (sensitive strain).
 The production of kappa particles is dependent on a dominant allele K, so that killer
strains are KKor Kk and sensitive strains are ordinarily kk. 
 In absence of dominant allele K, kappa particles cannot multiply and in absence of
kappa particles, dominant allele K cannot produce them de novo. 
 Consequently sensitive strains with genotypes KK or kk can be obtained. These will
not carry any kappa particles. However, killer strain with genotype kk cannot be
obtained, because even if kappa particles are present, these would be lost in absence of
dominant allele.
 If Paramecium clones with genotypes KK or Kk are allowed to multiply asexually at
such a fast rate, that division of kappa particles cannot keep pace with division of cells,
kappa particles will be eventually lost. Consequently sensitive strains with dominant
genotype (KK, Kk)having no kappa particles would be obtained.
If the killer (KK)and sensitive (kk)strains are allowed to
conjugate, all exconjugants (the cells separating after conjugation)
will have same genotype Kk. 
Phenotypes of these exconjugants will, however, depend upon
duration for which conjugation is allowed
If conjugation does not persist long enough for exchange of
cytoplasm, heterozygote (Kk) exconjugants will only have
parental phenotypes
 It means that killers will remain killers and sensitive will
remain sensitive even after conjugation .
If conjugation persists, sensitive strain will receive kappa
particles and will become killer, so that exconjugants will be killers
having genotype Kk 
MATERNAL EFFECT
 Maternal effect refers to an inheritance pattern
for certain nuclear genes in which the genotype of
the mother directly determines the phenotype of her
offspring
 Surprisingly, the genotypes of the father and offspring
themselves do not affect the phenotype of the offspring
 This phenomenon is due to the accumulation of
gene products that the mother provides to her
developing eggs
Shell coiling in snail
 The first example of a maternal effect
gene is seen morphological features
of the water snail, Lymnaea peregra
 In this species, the shell and internal
organs can be arranged in one of
two directions
 Right-handed (dextral)
 Left-handed (sinistral)
 The dextral orientation is more
common and dominant
 The snail’s body plan curvature
depends on the cleavage pattern
of the egg immediately after
fertilization
Many combinations of crosses produced results
that could not be explained by a Mendelian
pattern of inheritance Reciprocal cross Alfred Sturtevant
proposed the idea that
snail coiling is due to a
maternal effect gene
D is dominant to d (D or d allele)
(dextral is dominant to
sinistral)

A 3:1 phenotypic ratio would

be predicted by a Mendelian
pattern of inheritance

Pattern deviating from

Mendelian pattern
Reciprocal cross

F1 mothers are Dd

The dominant allele, D,

caused ALL the F2 offspring
to be dextral, even if the
offspring’s genotype was dd

This ratio corresponds to the

genotypes of the F2 females,
which were the mothers of F3
(F2 mothers include 3 with the D allele
and 1 with the d allele)

This explains this 3:1 ratio

 Thus, in this example
 DD or Dd mothers produce dextral offspring
 dd mothers produce sinistral offspring

 The phenotype of the progeny is determined by

the mother’s genotype NOT phenotype
 The genotypes of the father and offspring do not affect
the phenotype of the offspring
 A female is heterozygous for the snail-coiling
maternal effect gene
 The haploid oocyte received the d allele in meiosis
The Petite Trait in Yeast
 Mutations that yield defective mitochondria are expected
to make cells grow much more slowly

 Boris Ephrussi and his colleagues identified

Saccharomyces cerevisiae mutants that have such a
phenotype
 These were called petites because they formed small
colonies on agar plates
 Wild-type strains formed larger colonies
 Biochemical and physiological evidence indicated
that petite mutants had defective mitochondria

 Genetic analyses showed that petite mutants are

inherited in different ways
 Two main types of mutants were identified
 1. Segregational mutants
 Have mutations in genes located in the nucleus
 Segregate in a Mendelian manner in meiosis
 2. Vegetative mutants
 Have mutations in genes located in the mitochondrial genome
 Show a non-Mendelian pattern of inheritance
 Each resulting tetrad
Zygote then meiosis shows a 2:2 ratio of
wild-type to petite
Euphressi discovered two types of vegetative petites
Neutral and Suppressive

Zygote then meiosis Zygote then meiosis

Each resulting tetrad These results contradict the normal Each resulting tetrad
shows a 4:0 ratio of 2:2 ratio expected for the segregation shows a 0:4 ratio of
wild-type to petite of Mendelian traits wild-type to petite
 Researchers later found that

 Neutral petites lack most of their mitochondrial DNA

 Suppressive petites lack only small segments of

mtDNA

 When two yeast cells are mated, offspring inherit

mitochondria from both parents
  Progeny have both “wild type” and “suppressive
petite” mitochondria
 So how come only petite colonies are produced?
 Progeny have both “wild type”
 Two possibilities
and “neutral petite” mitochondria
 They display a normal  i. Suppressive petite mitochondria could
phenotype because of the wild replicate faster than wild-type mitochondria
type mitochondria  ii. Recombination between wild-type and
petite mtDNA may ultimately produce defects
in the wild-type mitochondria