Multiple Allele
Multiple Allele
In Mendelian inheritance a given locus of chromosome was occupied by 2 kinds of genes, i.e., a
normal gene (for round seed shape) and other its mutant recessive gene (wrinkled seed shape).
But it may be possible that normal gene may show still many mutations in pea besides the one
for wrinkledness. Here the locus will be occupied by normal allele and its two or more mutant
genes.
Thus, three or more kinds of genes occupying the same locus in individual chromosome are
referred to as multiple alleles. In short many alleles of a single gene are called multiple alleles.
The concept of multiple alleles is described under the term “multiple allelism”.
Dawson and Whitehouse in England proposed the term panallele for all the gene mutations at a
given locus in a chromosome. These differ from the multiple factor in one respect that multiple
factors occupy different loci while alleles occupy same locus.
“Three or more kinds of gene which occupy the same locus are referred to as
multiple alleles.” Altenburg
Characteristics of Multiple Alleles:
1. The study of multiple alleles may be done in population.
2. Multiple alleles are situated on homologous chromosomes at the same locus.
3. There is no crossing over between the members of multiple alleles. Crossing over takes place
between two different genes only (inter-generic recombination) and does not occur within a
gene (intragenic recombination).
5. Multiple alleles never show complementation with each other. By complementation test the
allelic and non-allelic genes may be differentiated well. The production of wild type phenotype
in a trans-heterozygote for 2 mutant alleles is known as complementation test.
6. The wild type (normal) allele is nearly always dominant while the other mutant alleles in the
series may show dominance or there may be an intermediate phenotypic effect.
7. When any two of the multiple alleles are crossed, the phenotype is of a mutant type and not
the wild type.
8. Further, F2 generations from such crosses show typical monohybrid ratio for the concerned
character.
Examples of Multiple Alleles:
1. Wings of Drosophila:
In Drosophila wings are normally long. There occurred two mutations at the same locus in
different flies, one causing vestigial (reduced) wings and other mutation causing antlered (less
developed) wings. Both vestigial and antlered are alleles of the same normal gene and also of
each other and are recessive to the normal gene.
Suppose vestigial is represented by the symbol ‘vg’ and antlered wing by ‘vg a‘. The normal allele
is represented by the symbol +.
Thus, there are three races of Drosophila:
(i) Long ++ (+/+)
When a fly with vestigial wing is crossed with another fly having antlered wings, the F 1 hybrids
are intermediate in wing length
showing that none of the mutated gene is dominant over the other. This hybrid is some times
said as the vestigial antlered compound and contains two mutated genes at the same locus. They
show Mendelian segregation and recombination.
Besides the vestigial and antlered wing described above there are several other mutations
occurring at the same locus and resulting in nicked wings, strap wings or no wings etc. These are
all multiple alleles.
Another example of multiple alleles is the eye colour in Drosophila. The normal colour of the eye
is red. Mutation changed this red eye colour to white. Other mutations at white locus took place
changing the red eye colour to various lighter shades like cherry, apricot, eosin, creamy, ivory,
blood etc., are also visible and are due to multiple alleles.
A cross between the two mutant forms, produces intermediate type in the F 1 except white and
apricot races which are not alleles but closely linked genes.
2. Coat Colour in Rabbit:
The colour of the skin in rabbits is influenced by a series of multiple alleles. The normal colour
of the skin is brown. Besides it there are white races called albino and Himalayan as the mutant
races. The Himalayan is similar to albino but has darker nose, ear, feet and tail. The mutant
genes albino (a) and Himalayan (ah) occupy the
same locus and are allelic. Both albino and Himalayan are recessive to their normal allele (+).
A cross between an albino and Himalayan produces a Himalayan in the F 1 and not intermediate
as is usual in the case of other multiple alleles.
3. Self-Sterility in Plants:
Kolreuter (1764) described self- sterility in tobacco (Nicotianalongiflora). The reason was done
by East. He described that self-sterility is due to series of alleles designated as s 1, s2, s3 and s4 etc.
The hybrids S1/S2 or S1/S3 or S3/S4 are self-sterile because pollen grains from these varieties did
not develop, but pollens of S1/S2 were effective and capable of fertilization with S3/S4.
The genes causing self-sterility in plants probably produce their effects by controlling the growth
rate of the pollen tubes. In compatible combinations, the pollen tube grows more and more
rapidly as it approaches the ovule, but in non-suitable ones, the growth of the pollen tube slows
down considerably, so that the flower withers away before fertilization can take place.
4. Blood Groups in Man:
Several genes in man produce multiple allelic series which affect an interesting and important
physiological characteristic of the human red blood cells. The red blood cells have special
antigens properties by which they respond to certain specific components (antibodies) of the
blood serum.
The antigen-antibody relationship is one of the great specificity like that between lock and key.
Each antigen and its associated antibody has a peculiear chemical configuration. Landsteiner
discovered in 1900 that when the red cells of one person are placed in the blood serum of
another person, the cells become clumped or agglutinated.
If blood transfusions were made between persons of two such incompatible blood groups, the
transfused cells were likely to clump and shut out the capillaries in the recipient, some times
resulting in death.
However, such reactions occurred only when the cells of certain individuals were placed in
serum from certain other persons. It was found that all persons could be classified in to four
groups with regard to the antigen property of the blood cells.
Large number of persons have been classified in to these four groups by means of the
agglutination test and the distribution of blood
groups in the offspring of parents of known blood groups has been studied. The evidence shows
that these blood properties are determined by a series of three allelic genes I A, IB and i, as
follows:
IA is a gene for the production of the anti-gin A. IB for antigen B, and i for neither antigen. The
existence of these alleles in man and the case with which the blood groups can be identified have
obvious practical applications in blood transfusion, cases of disputed percentage and description
of human populations.
The alleles of these genes which affect a variety of biochemical properties of the blood, act in
such a way that in the heterozygous
compound IAIB, each allele exhibits its own characteristics and specific effect. The cells of the
heterozygote contain both antigens A and B. On the other hand, I A and IB both show complete
dominance over i, which lacks both antigens.
Table showing possible blood types of children from parents of various blood groups.
The original discovery was that the red cells are agglutinated by a serum prepared by
immunizing rabbits against the blood of Rhesus monkey. The antigen responsible for this
reaction was consequently called as Rhesus factor and the gene that causes this property was
denoted as R-r or Rh-rh.
Interest in this factor was stimulated by Levine’s study of a characteristic form of anaemia,
known as Erythroblastosis foetalis, which occurs occasionally in new born infants.
It was found that the infants suffering from this anaemia are usually Rh-positive and so are their
fathers; but their mothers are Rh-negative. The origin of the disease was explained as follows:
The Rh+ foetus developing in the uterus of an Rh – mother causes the formation of mother’s
blood stream of anti Rh antibodies.
These antibodies, especially as a result of a succession of several Rh + pregnancies, gain sufficient
strength in the mother’s blood so that they may attack the red blood cells of the foetus. The
reaction between these antibodies of the mother and the red cells of her unborn child provokes
haemolysis and anaemia; this may be serious enough to cause the death of the newborn infant
or abortion of the foetus.
The blood stream of a mother who has had an erythroblastotic infant is a much more potent and
convenient reagent than sera of rabbits, immunized by blood of rhesus monkey’s for testing the
blood of other persons to distinguish Rh+ from Rh– individuals using such sera from woman who
had erythroblastotic infants, it was discovered that: there exist not one but several kinds of
Rh+ and Rh- persons. There are several different Rh antigens which are detected by specific
antisera.
Thus, an Rh– woman immunized during pregnancy by the Rh + children may have in her blood
serum antibodies, that agglutinate not only Rh+ red cells but also cells from a few persons
known to be Rh–.
By selective absorption two kinds of antibodies may be separated from such a serum, one known
as anti-D which agglutinates (= coagulates) only Rh + cells, the other known as anti-C which
agglutinates particular rare types of Rh–. Another specific antibody, known as anti-c agglutinates
all cells that lack C.
With these three antisera, six types of blood can be recognized. Studies of parent and children
show that persons of type Cc are heterozygous for an allele C determining C anti-gena. CC
persons are homozygous for C and cc are homozygous for c. There is obviously no dominance,
each allele producing its own antigen in the heterozygote as in the AB blood type.
No anti serum is available for detecting d, the alternative to D. D + persons may be heterozygous
or homozygous. However, the genotypes of such persons may be diagnosis from their progeny;
for example D+ person who has a d– child is thereby shown to be Dd.
Two other specific antibodies, anti-E and anti-c have been found. These detect the antigens E
and e determined by a pair of alleles E and e. The three elementary types of antigens C-c, D and
E-e, occur in fixed combinations that are always inherited together as alleles of a single gene.
Wiener and Fisher showed the existence of a series of eight different alternative arrangements of
these three types of Rh antigens and expressed them by means of following symbols.
There is not much difference between the two theories of Wiener and
Fisher. Wiener opinion is that there are multiple variations of one
gene whereas according to the view of Fisher three different genes
lying very close together are responsible for differences.
Pleiotropism:
The opposite of polygene effect is known as pleiotropism i.e., a single
gene influence or govern many characters.
For example, gene for vestigial wing influence the nature of halters
(modified balancers of Drosophila). The halters are not normal but
reduced in flies with vestigial wings. The vestigial gene also affects
position of dorsal bristles which instead of being horizontal turn out to
be vertical.
This gene also affects the shape of spermatheca i.e., the shape of
spermatheca is changed ; the number of egg strings in the ovaries is
decreased compared to normal when the vestigial larvae are well fed
but relatively increased when they are poorly fed ; length of life and
fruitfulness or fertility are lowered, and there are still other
differences.
Theories of Allelism:
Various theories have been put forward to explain the nature of
allelism origin and occurrence.
In maize the position effect are some times due to transposition (act of
changing place or order) of very minute particles of heterochromatin.
There are also sign or token that particles of different kinds of
heterochromatin suppress the expression of normal gene to different
degrees.
1. Pseudo alleles:
Alleles are different forms of the same gene located at the
corresponding loci or the same locus. Sometimes it has been found
that non-homologous genes which are situated at near but different
loci affect the same character in the same manner as if they are
different forms or alleles of the same gene. They are said as pseudo
alleles. These pseudo alleles which are closely linked show re-
combinations by crossing over unlike the alleles.
ADVERTISEMENTS:
3. Lsoalleles:
Sometimes, a dominant gene occurs in two or more forms. These
multiple dominant alleles will produce the same phenotypic effect in
homozygous condition but their effect will show a small difference in
heterozygous state.
In Drosophila, thus, the gene for red eye colour is dominant over
white. The red gene will produce dark red colour in the homozygous
condition but in combination with the white allele the gene for red
colour produces a dark red colour in flies from Soviet Russia but the
same combination in the flies coming from the U.S.A. produces a light
red colour. It does mean that dominant gene for red colour occurs in
two forms. These are said as isoalleles.
4. Phenocopy:
Characters are the result of interaction between the genotype and the
environment. When a gene mutates, its phenotypic effect also changes.
Some times, a change in the environment produces a visible change in
the phenotype of the normal gene which resembles the effect as
already known mutant.
The yellow colour of the endosperm in the yellow seeds is the result of
fertilization by pollen from the yellow variety. The yellow colour
indicates that the seeds are hybrids and the white seeds are
homozygous.
The yellow colour of the endosperm is dominant over white and when
the plants raised from the yellow seeds are self-pollinated, yellow and
white seeds are produced in the ratio of 3:1. Another example of xenia
may be exemplified. If a sweet corn (maize) is pollinated by a starchy
variety, the endosperm is starchy because the starchy gene introduced
by the pollen is dominant over its sugary allele.
6. Metaxenia:
It is the term used to describe the effect of foreign pollen on other
tissues belonging to the mother plant, outside the endosperm and
embryo. It is sometimes evident in the fruit and seed coats.
In the population, the coat colour gene has several alleles, and the pigment-
producing protein will be determined by the inheritance and expression of
these alleles. Curliness, shading, patterning, and even texture is all controlled
by the same genes. Because of the numerous potential genotype
combinations and expressions resulting from these genes, a broad range of
breeds exist. Even when two parents share just four alleles for each gene, the
variety may be amazing.
Cats’ coat colour phenotype — patches of black and orange (tortoiseshell
pattern), black, grey, white, and patches of white hair — strongly shows
multiple allelism, since there appear to be more than two alleles for the coat
colour phenotype (piebald spotting).
The genotype of a cat may usually be detected by looking at its coat colouring
and pattern. It is typically feasible to estimate the colouring possibilities of
kittens if the phenotypes of the parents are known, but the computations
would be difficult in most situations.
Pleiotropy
Pleiotropy is a situation in which more than one gene can affect the phenotype
in numerous ways. Pleiotropy entails more than one gene that influences
phenotype,
Whereas multiple allelism simply comprises a single gene with numerous
variations (referred to as many alleles).
Albino people, for example, are more likely than pigmented ones to have
crossed eyes. Albinos may also have a cross-eyed characteristic, in addition
to having insufficient colour production in their skin and hair. However, not all
albinos have this feature, showing that the two qualities are not connected in
these situations. (BioL110F2012 – Confluence, 2012).
The Complex Expression Patterns of Multiple Alleles Another example is the
colour of one’s eyes. More than one gene influences the characteristic. OCA2
and HERC2 are two of these genes.
Epistasis
When one gene influences the expression of another, this is known as
epistasis. When genes cooperate to generate a certain characteristic, this
happens.
One example is the determination of coat colour in some species (such as
horses), where the effect of one gene is influenced by the influence of another
gene that controls hair pigment deposition. (BioL110F2012 – Confluence,
2012). Multiple Alleles’ Complex Expression Patterns.