Sterility
Introduction
Male sterility refers
to the absence of functional pollen grains in other wise hermaphrodite flowers. Male
sterility is of great value in the production of hybrid seed. Male sterility is of two
types:
Genetic and
cytoplasmic. Cytoplasmic male sterility is termed cytoplasmic-genetic when restorer genes
are known. There are several natural mechanisms that control the mode of pollination in
crop plants. Two of these mechanisms, self-incompatibility and male sterility are of
special significance because of their utilization in hybrid seed production. Male
sterility is characterized by nonfunctional pollen grains, while female gametes function
normally.
Male sterility is
classified into three groups:
Genetic
Cytoplasmic
Cytoplasmicgenetic
Genetic male sterility
Genetic male sterility
is ordinarily governed by a single recessive gene, ms, but dominant genes
governing male sterility are also known, e.g., in safflower. Male sterility alleles arise
spontaneously or may be artificially induced. A male sterile line may be maintained by
crossing it with heterozygous male fertile plants. Such a mating produces 1:1 male sterile
and male fertile plants.
- Utilization in plant breeding
Genetic male sterility
may be used in hybrid seed production. The progeny from ms msxMs ms crosses are
used as female, and are interplanted with a homozygous male fertile (Ms Ms)
pollinator. The genotypes of the ms ms and Ms ms lines are identical except
for the ms locus, i.e., they are isogenic and are known as male sterile (A)
and maintainer (B) lines, respectively. The female line would, therefore, contain
both male sterile and male fertile plants; the latter must be identified and removed
before fertile plants; the latter must be identified and removed before pollen shedding.
Pollen dispersal from the male (pollinator) line should be good for a satisfactory seed
set in the female line. In India, it is being used for hybrid seed production of arhar
(C.catan) by some private seed companies. It is use in several other crops, e.g., cotton,
barley, tomato, sunflower, cucurbits etc., but it is not yet practically feasible.
Cytoplasmic Male Sterility
This type of male
sterility is determined by the cytoplasm. Since the cytoylasm of a zygote comes primarily
from egg cell, the progeny of such male sterile plants would always be male sterile. The
male sterile line is maintained by crossing it with the pollinator strain used as the
recurrent parent in the backcross programme since it s nuclear genotype is identical with
that of the male sterile line. Such a male fertile line is known as the maintainer line
or B line as it is used to maintain the male sterile line. The male sterile line is
also known as A line. It is observe in some important plant crops viz. Maize, (Zea
mays), Nicotiama tabacum, Triticum aestivum, Gossypium hrisutum, Sorghum bicolor,
Helianthus annus, Oryza sativa.
- Utilization in plant breeding
Cytoplasmic male
sterility may be utilized for producing hybrid seed in certain ornamental species, or in
species where a vegetative part is of economic value. But in those crop plants where seed
is the economic part, it is of no use because the hybrid progeny would be male sterile.
Cytoplasmic-Genetic male sterility
This is a case of
cytoplasmic male sterility where a nuclear gene for restoring fertility in the male
sterile line is known. The fertility restorer gene R, is dominant and is found in
certain strains of the species, or may be transferred from a related species e.g., in
wheat. This gene restores male fertility in the male sterile line, hence it is known as
restorer gene. The cases of cytoplasmic male sterility would be included in the
cytoplasmic genetic system as and when restorer genes for them would be discovered. This
system is known in maize, jowar, bajra, sunflower, rice and wheat.
The plants would be
male sterile in the presence of male sterile cytoplasm if the nuclear genotype were rr,
but would be male fertile if the nucleus were Rr or RR. The development of
new restorer strains is somewhat indirect. First a restorer strain (R) is crossed
with a male sterile line (A). the resulting male fertile plants are used as the
female parent in repeated backcrosses with the strain (C), used as the recurrent
parent, to which the transfer of restorer gene is desired. In each generation, male
sterile plants are discarded, and the male fertile plants are used as females for
backcrossing to the strain C. This acts as a selection device for the restorer gene
R during the backcross programme. At the end of the backcross programme, a restorer
line isogenic to the strain C would be recovered.
- Utilization in plant breeding
The
cytoplasmic-genetic male sterility is used commercially to produce hybrid seed in maize,
bajra and jowar. A triple cross may be produced by crossing single cross with a fertility
restoring inbred so that all the plants in the triple cross would be male fertile.
Origin of Male Sterile Cytoplasm
Mutant male sterile
cytoplasms arise spontaneously in low frequencies. Mutant cytoplasms have been isolated in
maize, bajra and sunflower.
- Interspecific hybridization
Transfer of the full
somatic chromo-some complement of a crop species, through repeated backcrossing into the
cytoplasm of a related wild species often leads to cytoplasmic male sterility. In
cross-pollinated crop species, the male sterile cytoplasms have generally originated
through mutation, while in self-pollinated crops they have been transferred from related
species.
- Induction through ethidium bromide
Ethidium bromide is a
potent mutagen for cytoplasmic genes or plasmagenes. Male sterile cytoplasm may be induced
by seed treatment with ethidium bromide e.g., Petunia.
Limitation of Cytoplasmic-Genetic Male Sterility
Undesirable - Effects of the Cytoplasm
for e.g.,
the Texas cytoplasm in maize, by far the most successful cytoplasm commercially, slightly
retards growth, yield (2.4%) plant height and leaf number; induces earlier silking and
delayed pollen shedding; and makes the plants highly susceptible to Helminthosporium leaf
blight. Restorer genes only restore male fertility; they are unable to remove the side
effects of the male sterile cytoplasms.
Unsatisfactory fertility restoration
In many
cases, restoration of fertility is not satisfactory and cannot be used in the production
of hybrid seed.
Unsatisfactory pollination
Natural
pollination is often not satisfactory, except in wind-pollinated crops like maize. This
reduces the production of hybrid seed, and thereby increases its cost. In some species
e.g., Capsicum, this has prevented the use of male sterility in hybrid seed
production. Poor pollination would always be a major problem in self-pollinators e.g.
wheat.
Modifier genes may reduce the
effectiveness of cytoplasmic male sterility.
Sometimes, cytoplasm may also be
contributed by the sperm which, in the long run, may lead to a breakdown of the male
sterility mechanism.
Male sterility mechanisms may break
down partially under certain environmental conditions resulting in some pollen production
by the male sterile lines.
In crops like wheat, polyploid nature
of the crop and undesirable linkages with the restorer gene make it very difficult to
develop a suitable restorer (R ) line.
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