Cytoplasmic Inheritance
Cytoplasmic Inheritance
Besides chromosomes, various organelles of cytoplasm also contain DNA. The mitochondria
and plastids have their own DNA and carry their genetic characters themselves. The mechanism in
which cytoplasmic inclusions (e.g., alpha, beta, sigma and kappa particles) and organelles (plastids,
mitochondria, centriole, etc) take part in transmission of characters from generation to generation is
called cytoplasmic inheritance. Since cytoplasmic inheritance is based on cytoplasmically DNA
molecules, it is also called extra chromosomal inheritance.
The smaller inheritable extra chromosomal unit is called as plasma gene and all the
plasmagenes of a cell constitute the Plasmon (like the genome).
Cytoplasmic inheritance is due to the plasmagenes located in cell organelles that are integral
constituents of normal cells. The characteristic features of this inheritance are summarized below.
In Mendelian inheritance, the results of reciprocal crosses are identical (one exceptional – sex
linked inheritance). If the character is transmitted through cytoplasm, the reciprocal cross results will
be different.
2. Somatic segregation
Plasma genes generally show somatic segregation during mitosis, a feature of rare occurrence
in the case of nuclear genes.
3. Non-mappability
Gene controlled characters shows linkages and hence they are mappable. But the characters
transmitted through cytoplasm show no linkage. Hence, they are not mappable.
4. Non-Segregation
There are two types of strains in Paramecium. One has kappa particles in its cytoplasm and
other does not have such particles in its cytoplasm and other does not have such particles. The
presence of kappa particles in the cytoplasm leads to production of a toxin known as paramecin. This
toxin can kill the strain Paramecium that lacks kappa particle. Thus the strain with kappa particle is
known as killer strain and that without kappa particle is called as sensitive strain.
The production of kappa particles is dependent on a dominant allele K, so that the killer
strains are KK or Kk and sensitive strains are ordinally kk. In the absence of dominant allele K, kappa
particles can not multiply and in the absence of kappa particles dominant allele k cannot produce them
de novo.
If the killer (KK) and sensitive (kk) strains are allowed to conjugate, all exconjugants (the
cells separating after conjugation) will have the same genotype Kk. The phenotypes of these
exconjugants will however depend upon duration for which conjugation is allowed. If conjugation
does not persist long enough for exchange for exchange of cytoplasm, heterozygote (Kk)
exconjugants will only have parental phenotypes. It means that killers will remain as killers and
sensitive will remain as sensitive after conjugation. If conjugation persists, sensitive stain will receive
kappa particles and will become killer, so that exconjugants will be killers having genotype Kk.
Plastid inheritance
Plastids are minute cytoplasmic organelles in plant cells. Most important are the
chloroplastids, which carry chlorophyll. Plastids arise from smaller cytoplasmic particles (plastid
primordial) that contain DNA. They duplicate themselves independently. They are transmitted
through the cytoplasm of the egg.
The Four-O’clock plant, mirabilis jalapa, has branches that produce either green, white or
mixed green-white (variegated) leaves. In crosses between flowers of these branches, the offspring are
all green if the maternal parent is a flower from a green branch. Such offspring remain green
throughout subsequent generations as long as maternal plant is green. Similarly, as long as the
maternal parent is from a white branch, the offspring are all white. When variegated branches are used
as female source, both green and plastids are present in cells of female parent. Therefore, female
gametes may carry either green or pale plastids or both. Consequently, three kinds of plants namely
green pale and variegated plants would be obtained.
Plasmids
Plasmids are called episomes. They are extra chromosomal, circular, covalently closed double
stranded DNA molecules found in bacteria. In effect, plasmids are accessory chromosomes. Plasmids
can replicate autonomously of the host chromosome. The size of plasmid ranges from two to several
hundred kilobases.
Plasmids carry genes for the inactivation of antibiotics, metabolism of natural products and
production of toxins. The F factors and R factors are important plasmids of Escherichia coli.
Mitochondria is present in living organisms arise from pre existing mitochondria. They are
small cytoplasmic organelles present in animal and plant cells but not present in bacteria and viruses.
Mictochondira provides cellular energy through oxidative phosphoriltion. Mitochondria contain a
small circular DNA molecule codes for limited number of structures and functions. The size of
mtDNA ranges form about 16 kb in mammals upto several hundred kilo base pairs in higher plants
(eg 570 kb in maize) and mt DNA usually found in multiple copies per organelle. The mtDNA play a
significant role in crop improvement. Recent evidences showed that the cytoplasmic genetic male
sterility system in crop plants is due to the interaction of mitochondrial genome to the nuclear
genome.
Chloroplast DNA
Chloroplast of the plant cell contain circular DNA molecule which are self-replicating in
nature. The isolated chloroplast found to capable of protein synthesis in the presence of light. The
DNA analysis revealed that 30-60 copies of the chloroplast genome are found in each chloroplast of
higher plants. The chloroplast genome contains herbicidal resistant and streptomycin resistant genes.
Maternal inheritance
Maternal effects are produced due to the influence of mother’s nuclear genotype on the
phenotype of its progeny and last for one generation. Characters showing the maternal effect exhibit
clear-cut differences in F1 for reciprocal crosses. One of the examples for maternal effect is coiling
pattern of shell in snail limnaea. In this snail, the direction of coiling of its shell is controlled by single
nuclear gene D/d; the dominant allele D produces right-handed coiling, while its recessive allele d
produces left-handed coiling. The direction of shell coiling in an individual is governed by the
genotype of its female parent and not by its own genotype. As a result, reciprocal crosses show
differences in coiling in F1 and there is no phenotypic segregation in F 2 the phenotypic effect of
segregation is observable in F3 only.
Crosses between females with left-handed coil (dd) and males having right handed coil (DD)
produce F1 progeny (Dd) with left-handed coil, since the genotype of the female parent is dd. In F 2
segregation of Dd produces three genotypes (DD,Dd,dd) in the ratio of 1:2:1. But the F 2 snails with
DD, Dd as well as dd genotypes exhibit right handed coiling since their female parent has the
genotype Dd which determines right-handed coiling in the progeny (irrespective of the genotypes of
the progeny). The F3 progeny from the F2 individuals with the genotypes DD and Dd will show right
handed coiling, while those from dd F 2 individuals will exhibit left-handed coiling of their shells; thus
produces the typical 3:1 ratio in F3.
The reciprocal cross (DDxdd), on the other hand, yields right-handed coiling in the F 1 (Dd) as
well as in the three genotypes, 1DD:2Dd:1dd, obtained in the F 2. But in F3 2/3 of the progenies show
right-handed coiling since they are derived from F 2 individuals having the genotypes DD and Dd. The
remaining 1/3 of the F3 progenies exhibit left handed coiling since their female parents had the
genotype dd; this yield the typical monohybrid ratio of 3:1 in the F3.
The direction of coiling in this snail is determined by the plane or the direction of the first
mitotic division of the zygote. The plane of the first division, of the other hand, is determined by some
substances already present in the egg cell. Obviously, these substances are produced by the female
parent; as a result, they would produce the phenotype appropriate for the maternal genotype. Further,
genotype of the zygote itself has no effect of the plane of first division and consequently, on the
direction coiling since its gene products are not involved in determining this trait. As a result, the
direction of coiling in an individual is governed by the genotype of its female parent. Therefore,
phenotypes appear one generation later than the appearance of the concerned genotypes, producing
delayed segregation in F3.
Iojap is characterized by contrasting stripes of green and white colours on leaves. This
character is controlled by a gene (y) located on chromosome 7 in corn. The dominant
allele Ij is responsible for normal green plastids and its recessive allele ij for iojap (striped)
character, so that Ijlj is green and ijij is iojap. Plants with genotype ijij would carry normal
green plastids as well as abnormal white plastids, while plants with genotype Ijij would carry
only green normal plastids. Female gametes from ijij can, therefore, carry either only green
plastids or only abnormal white plastids or both. Consequently, if ijij is used as female parent
and IjIj as male parent, the F1 individuals (Ijij)will be green, white or striped. From these
F1 individuals, ifIjij (striped) is again used as a female parent, and crossed withIjIj (green) as
male parent, two genotypes, namely Ijij and Ijijwould be obtained in equal proportions. Each
of these two genotypes would have three phenotypes (green, white and striped). It should be
noticed that, in such a cross, IjIj (striped) would also be obtained (Fig. 18.11).
It is, therefore, obvious that iojap character once inherited through female parent will be able
to maintain itself in the cytoplasm irrespective of genotype of plant. Therefore, as discussed
above, iojap or striped phenotype may have any of the three possible genotypes, IjIj,
Ijij and ijij.