MATERNAL INHERITENCE

If some treatments (chemical poison, heat shock etc.) are applied to the female parent, it may affect the egg’s

cytoplasm. As a result subsequent offspring’s are modified in some way. Effects of this kind are called Dauer-
modifications or persisting modifications.

It is observed that when protozoa are treated experimentally with chemical poisons or heat shocks, the treatments

induce several morphological abnormalities in them. Such abnormalities go on decreasing generation after generation
and, eventually, disappear completely through cell division if the treatments are removed.

Further evidences also come from fruit flies subjected to heat treatment and from bacteria treated with chemicals.

ii. Other kinds of maternal inheritance are also known which do not depend upon the repeated application of an

external stimulus to the cytoplasm. In this case, maternal inheritance is truly controlled by independent cytoplasmic
genes.

Maternal effects reflect the influence of the mother’s gene on developing tissues. Many important characteristics of
both animal and plants show maternal effects of which some examples axe described next.

(ii) Coiling of Snail Shells (Limnaea Peregra):

One of the earliest and classical examples of a maternal effect is that of the direction of coiling in shells of the water

snail Limnaea peregra. In this snail, the shell is spirally coiled. Usually the direction of coiling of the shell is clockwise

if viewed from the top of the shell. This type of coiling is called dextral. However, in some snails the coiling of shell is
anticlockwise. This type of coiling is sinistral.

The direction of shell coiling of both types of snail is governed by genotype of the female parent and not by their own
genotype. Further investigation suggests that coiling depends upon the early clearage in the zygote.

If the mitotic spindle is tilted to left (Fig. 22.1) of the median line of zygote, the successive cleavages will produce a

spiral to left (sinistral) and if the orientation of spindle is tilted to the right of the median line of zygote, the successive

cleavages will produce a spiral to right (dextral). The spindle orientation is controlled by the genotype of oocyte from
which the egg develops.
In a cross (Fig. 22.2) between dextral 9 (female) snail and sinistral (male), (follow the left side coloum of the Fig.

22.2) all the Fi progeny have dextral coils like their mother and also indicates that dextral character (RR) is dominant
over sinistral coiling (rr).

However, in the F1 x F1 cross (i.e., inbreeding or self fertilisation) all the F2 snails are also dextral. The F3 progeny from

F2 individuals with the genotype RR and Rr will show dextral coiling while those from rr F2 individual will exhibit

sinistral coiling of their shell; this produces the typical 3 : 1 ratio in F3 generation.

a) Chloroplast inheritance in variegated four ‘o’ clock plant.

The cytoplasmic or extra nuclear inheritance of colour in plant by plastids was first of all described by C. Correns in
1908 in the four o’clock plant, Mirabilis Jalapa. In contrast to other higher plants, Mirabilis contains three types of
leaves and parts: (1) Full green leaves or branches having chloroplast, (2) White (pale) leaves and branches having no
chloroplast, (3) Variegated branches having leukoplast in white (pale) areas and chloroplast in green patches (Fig.
47.3).

Because, the chlorophyll pigment of chloroplast is related with photosynthesis of food and leukoplasts are incapable
to perform photosynthesis, so the white or pale parts of plant survive by receiving nourishment from green parts.

Correns reported that flowers on green branches produced only green offsprings, regardless of the genotype and
phenotype of pollen parent and likewise, flowers from the white or pale branches produced only white or pale
seedings regardless of genotype and phenotype of pollen parent.

The plants developing from the white or pale seedings die because they lack chlorophyll and cannot carry on
photosynthesis. Correns further reported that flowers from the variegated branches yielded mixed progeny of green,
white (pale) and variegated plants in widely varying ratios (Fig. 47.4). These results are summarized in Table 47.1.
The irregularity of transmission from variegated branches could be understood by considering cytoplasmic genes
(plasmagenes) of plastids. A study of the egg during oogenesis in Mirabilis reveals that the ooplasm contains plastids
like cytoplasm of other plant cells.

If the egg cell is derived from green plant tissues, its ooplasm will contain coloured plastids; if derived from white
plant tissues, its ooplasm will contain white plastids; if derived from variegated tissues, its cytoplasm may contain
coloured plastids only, white plastids only or a mixture of coloured and white plastids. A study of the pollenogenesis,
however, reveals that pollen contains very little cytoplasm which in most cases is devoid of plastids. Without the
plastids, the pollen cannot affect this aspect of the offspring’s phenotype.

Mitotic segregation:
Variegated branches of Mirabilis Jalapa produce three kinds of eggs: some contain only white chloroplasts, some
contain only green chloroplasts and some contain both types of chloroplasts. In the subsequent mitotic divisions,
some form of cytoplasmic segregation occurs that segregate the chloroplast types into pure cell lines, thus, producing
the variegated phenotype in the progeny individual.

This process of sorting might be described as “mitotic segregation” of this is a pure extra nuclear phenomenon. In
mitotic segregation since both segregation and recombination of organelle genotype takes place, so it is called
cytoplasmic segregation and recombination (its acronym is CSAR).
In reciprocal cross (right side column of the Fig. 22.2) between dextral (male) and sinistral (female), all

the Fx progeny have sinistral coiling (Rr) instead of dextral coiling. In this case, F1 x F1 cross, all the F2 snails are, again,

dextral. This F3 progenies from F2 also exhibit the typical monohybrid ratio of 3 : 1.

(xiii) Uniparental Inheritance in Chlamydomonas Reinhardi:

R. Sager (1970) and N. Gilham (1968) have reported some cases of extra-chromosomal inheritance in green alga

Chlamydomonas reinhardi. The alga reproduces by asexual as well as sexual means. The sexual reproduction takes

place by fusion between two morphologically similar but physiologically dissimilar haploid gametes coming from
different haploid parents designated as ‘+’ and ‘-‘.
The gametic fusion produces the zygote. The sex is determined by a single chromosomal gene. When meiosis occurs in

the zygote, four haploid daughter protoplasts are formed which give rise to a new plant body. Although both the sexes
contribute equally to the zygote, there is maternal transmission of certain cytoplasmic traits.

Chlamydomonas is a haploid unicellulate green alga. It has two mating types—’+’ and ‘-‘.The two mating types are

governed by two alleles of a nuclear single gene. The

alleles axe named as mt+ and mt–. The + mating type is

considered as female, while the – mating type is

regarded as male. During sexual reproduction one

mt+ and one mt–cell pair and fuse together to form a

zygote where there is mixture of cytoplasm coming from

both mt+ and mt– gametes.

The zygote undergoes meiosis to produce 4 haploid

meiozoospores of which two zoospores contain ‘+’

alleles and other two contain ‘-‘ alleles, i.e., it shows

typical 1 : 1 segregation for nuclear genes. But for

their plasma genes all zoospores are identical and

contain only mt+type plasma genes by mt+ plasma

genes. The inactivation is not clear but it may involve

an enzymatic process.

R. Sager isolated two strains of Chlamydomonas: one

strain was resistant (Sr) to 500/xg of streptomycin per

ml of culture solution and the other one is sensitive. The

trait of streptomycin resistance is believed to be located

in its cp-DNA (chloroplast DNA).

Mating between mt+ streptomycin resistant (Sr) and mt sensitive (Ss) cells produce only resistant progeny but the

nuclear genes for mating type segregate as expected (Fig. 22.10). But the reciprocal cross between mt+ susceptible

and mt– resistant shows again the expected segregation for mating type but all progenies are sensitive type. Therefore,

it clearly provides an example for extra-nuclear inheritance. It is also observed that in less than 0.1% of zygotes

plasma genes from mt” parent are not inactivated and produce cytohets, i.e., heterozygotes for cytoplasmic genes.
mt+ sm-r × mt– sm-s → progeny all sm-r
mt+ sm-s × mt– sm-r → progeny all sm-s
Here, occurs a difference in reciprocal crosses; all progeny cell show the streptomycin phenotype of the mt + parent.
Like the maternal inheritance this is a case of uniparental inheritance. In fact, Sagar now refers to the mt+ mating type
as the female, using this analogy.
(ii) Poky strain of Neurospora:
In fungi, Neurospora crassa a number of mutations of mitochondria are inherited via the female parent. The best
studied of these is the poky strain of N. crassa, first isolated by Mitchell and Mitchell (1952). A poky mutant differs
from wild type strain of Neurospora in the following aspects: (1) it is slow growing; (2) it shows maternal inheritance,
and (3) it has abnormal cytochromes. Of the three cytochromes—cyt a, b and c found in wild type, cyt a and cyt b are
absent, and cyt c is in excess in poky mutant. In reciprocal crosses, poky character shows maternal inheritance:

Poky (female) × wild type (male) → all poky

Wild type (female) × poky (male) → all wild type

However, there are other marker nuclear genes (ad+/ad–) which show 1:1 Mendelian segregation. The following
evidences suggested that poky trait may be located in mitochondrial DNA: (i) slow growth may be due to lack of ATP
energy and source of this energy is mitochondria; (ii) cytochromes in poky strain differ from those in wild type in
quality and quantity and these cytochromes are found in mitochondria.

Extra-Nuclear Inheritance by Mitochondria of Yeast:

Yeast, Saccharomyces cerevisiae, are unicellular ascomycetes fungi. In this fungi, sexual reproduction takes place by

the fusion of two somatic cells to form a diploid zygote nucleus. Next follows two successive nuclear divisions forming

four haploid daughter nuclei, all of which take part in ascospore formation. Now the mother cell, i.e., zygote cell, is
called ascus.

The diploid zygote can also be grown vegetatively as a diploid strain that will later sporulate. Respiration of yeast cell

takes place both aerobically and anaerobically (fermentation). Certain mutant yeast cells are unable to utilise oxygen

and are comparatively small- sized and slow growing producing small colonies on agar medium. These small colonies
forming mutant strains of yeast are known as petites.

In petite strains, the necessary components (cytochrome b, c1) and some enzymes (cytochrome oxidase a, a3) for

aerobic terminal respiration activity are absent. But these components are present in the cell of normal strain where

they are associated with the inner membrane of mitochondria.

Petite strain can be maintained indefinitely in the vegetative state and can be mated with normal yeast cells. When
such mating are carried out, three petite varieties can be classified:
Nuclear (segregational) Petites:
When a normal haploid strain of yeast is crossed with a haploid petite strain, a normal diploid zygote is produced. The

haploid ascospores produced from zygote by sporulation are segregated in the ratio 1 : 1 (petite : normal). Hence the
result of such cross follows ordinary nuclear mendelian inheritance (Fig. 22.5).

Neutral Petites:

In this type, only normal wild ascospores are produced from mattings between petite and normal strain of yeast. The

petite characteristics is absent in the product of segregation. So it shows the non-Mendelian inheritance. This non-

Mendelian behaviour is very difficult to explain on the basis of nuclear genes and indicates that such petite
characteristics are caused by extra-nuclear inheritance.

Suppressive Petites:

In this type, all ascospores produced from mating between normal and petite strain are petite type. Such petites seem

to suppress normal respiratory behaviour and the suppressive petite factor acts as a dominant. Fig. 22.5 shows
diagrammatic scheme for explaining some differences between neutral and suppressive petite in terms of DNA.

Therefore, there are two different genetic causes for respiratory deficiency in yeast. One is nuclear and the other is

extra nuclear. On this basis a neutral petite having the nuclear gene for normally functioning mitochondria is crossed

with a segregational petite (Fig. 22,6). The diploid zygote produced from such cross can use the normal nuclear genes
from neutral petite and respire normally.
When such diploid zygotes are grown vegetatively as a diploid strain, they produce diploid colonies that are of normal

size and respire normally. But when such diploid zygotes are allowed to sporulate, they undergo meiosis and produce
four haploid ascospores of which two are petites and other two are normal.

It indicates that normal and petite characteristics segregate in the 1 : 1 ratios expected from mendelian segregation.
It is noted that the neutral petite contains the normal nuclear gene for the respiratory enzymes but the segregational

petite does not contain respiratory enzymes, so it is obvious that the cytoplasmic factor of the neutral petite appears
in the cytoplasm of diploid zygotes where the factor is possibly independent of nuclear control.

It is also noted that neutral petite strains are readily produced by subjecting normal strain in low doses of acriflavines

dyes as well as ethidium bromide. But the treatment of such doses of dye does not induce any nuclear changes. Thus it

strongly indicates the involvement of extra-nuclear change of gene controlling petite characteristics. Such changes
ultimately shows the extra-nuclear inheritance of petite characteristics.