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Gene Interactions
Gene Interactions
Genes decide the genotype and phenotype of an organism. Sometimes, the effects of
one gene gets modified by the presence or absence of various other genes.
In such a process, two or more types of genes can act simultaneously to modify the
expression of the other gene. The expression of one gene depends on the presence or
absence of another gene in any organism.
Explanation
Mendel’s dihybrid cross:
We know that Mendel simultaneously examined two different genes that controlled two
different characters in his dihybrid cross with pea plant.
For example, in one series of his experiments, Mendel began by crossing a plant that
was homozygous for the traits round seed shape and yellow seed color (RRYY), with a
different plant that was homozygous for the traits wrinkled seed shape and green seed
color (rryy).
When Mendel crossed the F1 progeny of the two plants with each (RrYr x RrYy), he
obtained an F2 generation with a phenotypic ratio of 9: 3: 3: 1.
In this dihybrid cross, each gene locus has an independent effect on a single
phenotype. Thus, the R and r alleles affected only the shape of the seed and had no
influence on the seed color, while the Y and y alleles only affected the seed color and
had no influence on the seed shape.
In this case, there are two separate genes that coded for two separate characteristics.
But imagine what will happen if two different loci affect the same characteristics?
What if, in Mendel’s experiment, both loci had an effect on seed color?
When two genes are involved in the expression of one characteristic, a dihybrid cross
can produce a phenotypic ratio that is very different from 9: 3: 3: 1.
Under such conditions, there are more than two gene products that affect the same
phenotype, and these products may have complex interactions.
It is the influence of alleles and non-alleles on the normal phenotypic expression of the
genes.
Incomplete dominance:
We have already studied that incomplete dominance is a phenomenon in which a
dominant allele or a form of a gene does not entirely hide or mask the effects of a
recessive allele, and the organism’s resulting phenotype shows a blending of both
alleles.
The homozygotic forms (𝐼𝐴 𝐼𝐴 and 𝐼𝐵 𝐼𝐵 ) express either the A or the B phenotype, and
heterozygotes (𝐼𝐴 𝐼𝐵) express both phenotypes equally.
The 𝐼𝐴 𝐼𝐵 individual has blood type AB. In a self-cross between heterozygotes expressing
a co-dominant trait, the three possible offspring genotypes are phenotypically distinct.
In this case, the F2 genotype and phenotype ratio are the same, i.e., 1: 2: 1. Thus, the
mendelian monohybrid ratio of 3: 1 gets modified.
Multiple alleles:
Individuals usually have only two alleles for each gene. Within a population, however,
there may be three or more alleles for a single gene trait. Such a trait is termed multi-
allelic.
For example, in humans, the ABO blood group is a good example of a single gene that
has multiple alleles.
Blood cells have molecular markers on their surfaces, and these play an important role
in allowing a person’s own body cells to be recognized by the immune system as ‘self ’.
To represent multi-allelic blood groups using correct genetic notation, the gene is
denoted as I and the three alleles are represented by superscripts: 𝐼^𝐴, 𝐼^𝐵 and 𝐼^𝑖.
Alleles A and B are codominant, as they each produce a molecular marker on red blood
cells. If both alleles are present, the blood cells have both markers. The i allele
produces no molecular marker on the red blood cells and is recessive to both A and B.
As a result, there are four possible phenotypes for the ABO blood system: a person may
have blood group A, B, AB or O. There are, however, six possible genotypes.
Another example of a gene that has multiple alleles is the gene for coat color in rabbits.
There are four alleles, called normal, chinchilla, Himalayan and albino.
In rabbits, there are four alleles which code for coat color: C, 𝑐h, 𝑐h and c. Allele C is
dominant to all the other alleles and produces a full color coat.
Allele c is recessive and this leads to an albino phenotype when the genotype is
homozygous recessive.
Full color coat is dominant over chinchilla, which is dominant over Himalayan, which is
dominant over albino.
Multiple alleles increase the number of genotypes and phenotypes that can be
produced. Without multiple-allele dominance, two alleles, such as T and t, create only
three possible genotypes—in this example, TT, Tt, and tt—and two possible
phenotypes.
However, the four alleles for rabbit-coat color produce ten possible genotypes and four
phenotypes. More variation in rabbit coat color comes from the interaction of the color
gene with other genes, such as the agouti gene or the broken gene.
Summary
Gene is defined as the basic unit of heredity.
Genes decide the genotype and phenotype of an organism. Sometimes, the effects of
one gene gets modified by the presence or absence of various other genes.
The interaction of genes present at different loci that affect the expression of the same
character is known as gene interaction. It is the influence of alleles and non-alleles on
the normal phenotypic expression of the genes.
Gene interactions are of two types:
1. Inter-allelic or intra-genic gene interaction:
In such cases, two alleles located on the same gene locus on two homologous
chromosomes of a gene interact to produce phenotypic expression.
Example: co-dominance, incomplete dominance, multiple alleles.
Incomplete dominance is a phenomenon in which a dominant allele or a form of a gene
does not entirely hide or mask the effects of a recessive allele, and the organism’s
resulting phenotype shows a blending of both alleles.
Co-dominance is a mode of inheritance in which both the alleles of a gene pair in a
heterozygote are fully expressed. As a result, the phenotype of the offspring is a
combination of the phenotype of the parents.
Individuals usually have only two alleles for each gene. Within a population, however,
there may be three or more alleles for a single gene trait. This is termed as multiple
allelism.
The second type of gene interactions is:
Gene Interactions: Meaning and Characteristics | Genetics (biologydiscussion.com)
https://www.biologydiscussion.com/genetics/gene-interactions/gene-interactions-meaning-and-
characteristics-genetics/37790
The term epistasis was first used by Bateson in 1909 to describe two
different genes which affect the same character, one of which masks
the expression of other gene. The gene that masks another gene is
called epistatic gene, and the gene whose expression is masked is
termed as hypostatic gene. Epistasis is also referred to as inter-genic
or inter-allelic gene interaction.
iii. Expression:
The phenotypic expression of one gene usually depends on the presence or absence of
epistatic gene. The gene which has masking effect is called epistatic gene and the gene
whose effect is masked is known as hypostatic gene.
When a cross was made between rose (RRpp) and pea (rrPP), a new phenotype called
walnut developed in F1. The walnut comb developed as a consequence of combining two
dominant alleles R and P together in F1. Inter-mating of F1 birds produced four types of
combs, viz., walnut, rose pea and single in 9 : 3 : 3 : 1 ratio in F 2 generation.
Here individuals with R-P-(9/16) genotypes produce walnut comb, because two dominant
genes together produce walnut comb. Individuals with R-pp (3/16) will give rise to rose
comb, and those with rrP-(3/16) genotypes will produce pea comb. The single comb (1/16)
will develop from a double recessive, genotype (Fig. 8.1).
Gene Interactions: Allelic and Non-Allelic | Cell Biology (biologydiscussion.com)
https://www.biologydiscussion.com/genetics/gene-interactions/gene-interactions-allelic-and-non-
allelic-cell-biology/38795
Contents:
1. Introduction to Gene Interaction
2. Allelic Gene Interactions
3. Non Allelic Gene Interactions
4. Multiple Factors and Polygenic Gene Interactions
5. Other Kinds of Gene Interactions
1. Introduction to Gene Interaction: Mendelian genetics does not explain all kinds of
inheritance for which the phenotypic ratios in some cases are different from Mendelian
ratios (3:1 for monohybrid, 9:3:3:1 for di-hybrid in F2). This is because sometimes a
particular allele may be partially or equally dominant to the other or due to existence of
more than two alleles or due to lethal alleles. These kinds of genetic interactions between
the alleles of a single gene are referred to as allelic or intra- allelic interactions.
Non-allelic or inter-allelic interactions also occur where the development of single character
is due to two or more genes affecting the expression of each other in various ways.
Thus, the expression of gene is not independent of each other and dependent on the
presence or absence of other gene or genes; These kinds of deviations from Mendelian one
gene-one trait concept is known as Factor Hypothesis or Interaction of Genes (Table 7.1).
2. Allelic Gene Interactions:
ADVERTISEMENTS:
In snapdragon and Mirabilis jalapa, the cross between pure bred red-flowered and white-
flowered plants yields pink-flowered F1 hybrid plants (deviation from parental phenotypes),
i.e., intermediate of the two parents. When F1 plants are self-fertilized, the F2 progeny shows
three classes of plants in the ratio 1 red: 2 pink: 1 white instead of 3:1 (Fig. 7.1).
Therefore, a F1 di-hybrid showing incomplete dominance for both the characters will
segregate in F2 into (1 :2 : 1) X (1 :2 : 1 ) = 1 :2 : 1 : 2 : 4 : 2:1 : 2 : 1. And a F 1 di-hybrid
showing complete dominance for one trait and incomplete dominance of another trait will
segregate in F2 into (3:1) x (1 :2:1) = 3:6:3:1 :2:1.
Co-dominance:
Here both the alleles of a gene express themselves in the heterozygotes. Phenotypes of both
the parents appear in F1 hybrid rather than the intermediate phenotype. In human, MN
blood group is controlled by a single gene.
Only two alleles exist, M and N. Father with N blood group (genotype NN) and mother with
M blood group (genotype MM) will have children with MN blood group (genotype MN).
Both phenotypes are identifiable in the hybrid. F 2 segregates in the ratio 1M blood group: 2
MN blood group : 1 N blood group.
ADVERTISEMENTS:
Over-dominance:
Sometimes the phenotype of F1 heterozygote is more extreme than that of either parents.
The amount of fluorescent eye pigment in heterozygous white eyes of Drosophila exceeds
that found in either parents.
Lethal Factor (2:1):
The genes which cause the death of the individual carrying it, is called lethal factor.
Recessive lethals are expressed only when they are in homozygous state and the
heterozygotes remain unaffected. There are genes that have a dominant phenotypic effect
but are recessive lethal, e.g., gene for yellow coat colour in mice.
ADVERTISEMENTS:
But many genes are recessive both in their phenotypic as well as lethal effects, e.g., gene
producing albino seedlings in barley (Fig. 7.2).
Dominant lethals are lost from the population because they cause death of the organism
even in a heterozygous state, e.g., epiloia gene in human beings. Conditional lethals require
a specific condition for their lethal action, e.g., temperature sensitive mutant of barley
(lethal effect at low temperature).
Balanced lethals are all heterozygous for the lethal genes; both dominant and recessive
homozygotes will die, e.g., balanced lethal system in Oenothera. Gametic lethals make the
gametes incapable of fertilization, e.g., segregation distorter gene in male Drosophila.
ADVERTISEMENTS:
Semi-lethal genes do not cause the death of all the individuals, e.g., xentha mutants in some
plants.
Multiple Alleles:
A gene for particular character may have more than two allelomorphs or alleles occupying
same locus of the chromosome (only two of them present in a diploid organism). These
allelomorphs make a series of multiple alleles.
Human ABO blood group system furnishes best example. The gene for antigen may occur in
three possible allelic forms – lA, IB, i. The allele for the A antigen is co-dominant with the
allele I8 for the B antigen. Both are completely dominant to the allele i which fails to specify
any detectable antigenic structure. Therefore, the possible genotypes of the four blood
groups are shown in Fig. 7.3.
Self-sterility in tobacco is determined by the gene with many different allelic forms. If there
are only three alleles (s1, s2, s3), the possible genotypes of plants are s 1s2, s1s3, s2s3 (always
heterozygous), homozygous genotypes (s1s1, s2s2, s3s3) are not possible in a self-incompatible
species.
In this case, pollen carrying an allele different from the two alleles present in the female
plant will be able to function resulting in restriction of fertility (Fig 7.4).Iso-alleles express
themselves within the same phenotypic range, e.g., in Drosophila several alleles (W +s ,W+c,
W+g) exhibit red eye colour.
ADVERTISEMENTS:
The inheritance of comb types in fowls is the best example where R gene gives rise to rose
comb and P gene gives rise to pea comb; both are dominant over single comb; the presence
of both the dominant genes results in walnut comb (Fig. 7.5). Similar pattern of inheritance
is found in Streptocarpus flower colour (Fig. 7.6).
Complementary Factor (9:7):
Certain characters are produced by the interaction between two or more genes occupying
different loci inherited from different parents. These genes are complementary to one
another, i.e., if present alone they remain unexpressed, only when they are brought together
through suitable crossing will express.
In sweet pea (Lathyrus odoratus), both the genes C and P are required to synthesize
anthocyanin pigment causing purple colour. But absence of any one cannot produce
anthocyanin causing white flower. So C and P are complementary to each other for
anthocyanin formation (Fig. 7.7).
Involvement of more than two complementary genes is possible, e.g., three complementary
genes governing aleurone colour in maize.
Epistasis:
When a gene or gene pair masks or prevents the expression of other non-allelic gene, called
epistasis. The gene which produces the effect called epistatic gene and the gene whose
expression is suppressed called hypostatic gene.
Dominant gene C produces black colour, absence of it causes albino. Gene A produces
agouti colour in presence of C, but cannot express in absence of it (with cc) resulting in
albino. Thus recessive allele c (cc) is epistatic to dominant allele A (Fig. 7.8).
The grain colour in maize is governed by two genes — R (red) and Pr (purple). The recessive
allele rr is epistatic to gene Pr (Fig. 7.9).
(b) Dominant Epistasis (12:3:1):
Sometimes a dominant gene does not allow the expression of other non-allelic gene called
dominant epistasis. In summer squash, the fruit colour is governed by two genes. The
dominant gene W for white colour, suppresses the expression of the gene Y which controls
yellow colour. So yellow colour appears only in absence of W. Thus W is epistatic to Y. In
absence of both W and Y, green colour develops (Fig. 7.10).
Inhibitory Factor:
Inhibitory factor is such a gene which itself has no phenotypic effect but inhibits the expres-
sion of another non-allelic gene; in rice, purple leaf colour is due to gene P, and p causing
green colour. Another non-allelic dominant gene I inhibits the expression of P but is
ineffective in recessive form (ii). Thus the factor I has no visible effect of its own but inhibits
the colour expression of P (Fig. 7.11).
Gene A or B alone gives rise to awns of medium length (the effect of A is same as B); but
when both present, long awn is produced; absence of both results aweless (Fig. 7.13).
Duplicate Gene (15:1):
Sometimes a character is controlled by two non-allelic genes whose dominant alleles
produce the same phenotype whether they are alone or together. In Shepherd’s purse
(Capsella bursa-pastoris), the presence of either gene A or gene B or both results in
triangular capsules; when both these genes are in recessive forms, the oval capsules
produced (Fig. 7.14).
Duplicate Gene with Dominance Modification (11:5):
A character controlled by two gene pairs showing dominance only if two dominant alleles
are present. Dominant phenotype will thus be produced only when two non-allelic
dominant alleles or two allelic dominant alleles are present. Such a case is found in pigment
glands of cotton (Fig. 7.15).
The hypothesis states that for a given quantitative trait there could be several genes, which
are independent in their segregation and had cumulative effect on the phenotype.
Kernel colour in wheat is a quantitative character and controlled by two different genes. The
heterozygote is intermediate in colour between the two homozygotes. Both the dominant
genes have small and equal (or almost equal) effects on seed colour. F 1 heterozygote for two
genes will segregate in F2 in the ratio 1:4: 6:4:1.
The intensity of seed colour depends on the number of dominant alleles present, i.e., their
effects are cumulative in nature (Fig. 7.16). It is now known that there are three genes
involved in kernel colour in wheat, thus a F, heterozygous for all three genes will segregate
in F1 in the ratio 1 : 6 : 15 : 20 : 15 : 6 : 1.
Corolla length in Nicotiana is another quantitative character. The crosses made between two
inbred varieties of N. longiflora or N. tabaccum (tobacco) differing in corolla length, show
F1 with uniform corolla length but F2 exhibits a greater degree of variation.
Mean value of F3 derived from single F2 plant with particular corolla length differs greatly
from other single plant F3 progenies. It is thus obvious that F2 plants differ genetically (Fig.
7.17).
Skin colour in human beings is under polygenic effect, the number of gene pairs involved
may be two (Fig. 7.18) or more than two, possibly four or five.
The number of genes involved in polygenic inheritance can be calculated from the frequency
of parental type using the formula 1/4n (n = number of gene pairs).
If parental type obtained is one out of every 64 offspring’s ( 1/64), then the number of genes
involved will be three (4n = 64 = 43).
ADVERTISEMENTS:
Suppressors:
Genes which will not allow mutant allele of another gene to express resulting in wild
phenotype called suppressor gene, e.g., Su-s in Drosophila suppresses the expression of
dominant mutant gene star eye(s).
Pleiotropy:
Gene having more than one effect (multiple effects) are called pleiotropic genes. They have a
major effect in addition to secondary effect. In Drosophila, the genes for bristle, eye and
wing significantly influence the number of facets in bar-eyed individuals.
Atavism:
The appearance of offspring’s which resemble their remote ancestors called atavism.
Penetrance:
The ability of a gene to be expressed phenotypically to any degree is called penetrance.
Penetrance may be complete, e.g., in pea, expression of R allele for red flower in
homozygous and heterozygous conditions. It may be incomplete, e.g., dominant gene P for
Polydactyly in human, sometimes does not show polydactylous condition in heterozygous
state.
Expressivity:
A trait though penetrant, may be quite variable in its phenotypic expression, e.g., in human
polydactylous condition may be penetrant in left hand but not in the right.
Gene Interactions
The genes of an individual do not operate isolated from one another, but
obviously are functioning in a common cellular environment. Thus, it is
expected interactions between genes would occur. Bateson and Punnett
performed a classical experiment that demonstrated genetic interactions.
They analyzed the three comb types of chicken known to exist at that time:
Rose Pea
Single Walnut
Result: The F1 differed from both parents and two new phenotypes not seen in
the parents appeared in the F2. How can this result be explained? The first
clue is the F2 ratio. We have seen this ratio before when the F1 from a dihybrid
cross is selfed (or intermated). This observation suggests that two genes may
control the phenotype of the comb. The gene interactions and genotypes were
determined by performing the appropriate testcrosses.
It was later shown that the genotypes of the initial parents were:
Rose = RRpp
Pea = rrPP
The development of any individual is obviously the expression of all the genes
that are a part of its genetic makeup. Therefore, it is not an unexpected
conclusion that more than one gene could be responsible for the expression
of a single phenotype. We will now discuss this situation. First let's give a
definition.
The interactions of the two genes which control comb type was revealed
because we could identify and recognize the 9:3:3:1. Other genetic
interactions were identified because the results of crossing two dihybrids
produced a modified Mendelian ratio. All of the results are modifications of the
9:3:3:1 ratio.
Thus, if a pure line wheat plant with a colored kernel (genotype = AABB) is
crossed to plant with white kernels (genotype = aabb) and the resulting
F1 plants are selfed, a modification of the dihybrid 9:3:3:1 ratio will be
produced. The following table provides a biochemical explanation for the 15:1
ratio.
If we sum the three different genotypes that will produce a colored kernel we
can see that we can achieve a 15:1 ratio. Because either of the genes can
provide the wild type phenotype, this interaction is called duplicate gene
action.
If two genes are involved in a specific pathway and functional products from
both are required for expression, then one recessive allelic pair at either allelic
pair would result in the mutant phenotype. This is graphically shown in the
following diagram.
If a pure line pea plant with colored flowers (genotype = CCPP) is crossed to
pure line, homozygous recessive plant with white flowers, the F1 plant will
have colored flowers and a CcPp genotype. The normal ratio from selfing
dihybrid is 9:3:3:1, but epistatic interactions of the C and P genes will give a
modified 9:7 ratio. The following table describes the interactions for each
genotype and how the ratio occurs.
Because both genes are required for the correct phenotype, this epistatic
interaction is called complementary gene action.
With this interaction, color is recessive to no color at one allelic pair. This
recessive allele must be expressed before the specific color allele at a second
locus is expressed. At the first gene white colored squash is dominant to
colored squash, and the gene symbols are W=white and w=colored. At the
second gene yellow is dominant to green, and the symbols used
are G=yellow, g=green. If the dihybrid is selfed, three phenotypes are
produced in a 12:3:1 ratio. The following table explains how this ratio is
obtained.
Shapes of Squash Fruit