Interactions of genes

Gene interactions
A single character can be governed by two or more genes. They are called non - allelic or
intergenic genetic interactions. The independent genes (non-homologous) located on the same
or on different chromosomes interact with one another for the expression of a single
phenotypic trait of an organism.
Types of genetic interactions
The gene is a chemical determiner. The phenotypic trait results from the combined action of
many genes and their products constantly interacting with the environment.
Phenotype=Genotype + Environment
 The environment includes not only ecological factors such as temperature and light,
but also internal factors such as hormones and enzymes.
 The enzymes are proteins and the specific molecular organization of protein is
determined by genes.
 Biochemical studies indicated there is precursor and end products. A simplest
biosynthetic pathway includes various steps, each step is catalyzed by a specific enzymatic
protein and each enzymatic protein in its turn depends on a specific gene for its production.
Gene D produces enzyme D that act on IP 3(Intermediate Product) that produces an end
product. This gives the final phenotype.
 If there is mutation in gene D, there is a break in the biosynthetic pathway. There will
not change from IP3 to end product.
 A different end product may be produced called as metabolic block. This is caused
because of the mutation occurred in the particular gene.
 Here two or more genes interact together to produce a particular phenotype.
Epistasis
 A gene or locus which suppressed or masked the action of a gene at another locus was
termed suppressor orepistatic gene.
 The gene or locus which was suppressed by an epistatic gene was called hypostatic
gene.
 Dominance is the masking effect which one allele has upon the expression of another
allele at the same locus (i.e, intra-genic or inter allelic gene suppression).

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  Epistasis involves inter-genic suppression or the masking effect which one gene
locus has upon the expression of another. Thus, epistasis refers to variation resulting from the
interaction of alleles at different loci.
 Epistatic interaction between two loci produces variation on the 9:3:3:1 F2 phenotypic
ratio that occur when there are two dominant genes which produce different phenotypes.
 When there is epistasis, the number of F 2 phenotypes is usually reduced from four to
either two or three,depending on the type of epistasis.
Table 5.1. Autosomal phenotypes controlled by epistasis

Species Genes Phenotypes

Common carp S, N Scale pattern
B1 , B2 Orange body colour
Gold fish M, S Albino
DP1 , DP2 Depigmentation of melanophores

Body colour in many tropical fish is controlled by epistatic interactions between or among
two or more loci. A set of qualitative phenotypes may be controlled by more than two genes.
 Body colour in the Siamese fighting fish is an example of a set of phenotypes that is
controlled by the epistatic interaction among four genes.
 Working with these phenotypes is far more complicated because of the number of genes
involved. Fortunately,in food fish, no qualitative phenotype controlled by more than two
genes has been discovered.
 Scale pattern in common carp is the most important phenotype controlled by epistasis. An
understanding of the inheritance of scale pattern in common carp is of great important because
common carp with a reduced scale pattern has a higher market price in Europe. Common wild
type scale pattern (scaled) is more desired in Asia. Since common carp forms the world’s
most important cultured food fish, the genetic technique to produce the desired scale
phenotype can be of tremendous importance. In growth rate, scaled carp are better than the
scattered ones and the linear ones are better than the nude ones.
 Scale pattern in common carp is controlled by the S and the N genes. They produce the
phenotypes through a type of dominant epistasis where the N gene is the epistatic locus, but it

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 is a dominant lethal epistatic gene (it exhibits incomplete dominance and is lethal with
homozygous state).
 The S gene controls the scaliness, and the N gene modifies the pattern. The S allele is
completely dominant over thes allele. In the presence of the gene N the action of the gene S is
stronger. S allele is the common wild type, s allele produces reduced number of scales, and
those that remain are greatly enlarged (mirror).
 A single N allele changes scaled carp into line carp (scales limited to the dorsal and ventral
margins and the lateral line) and changes mirror common carp into leather or nude common
carp (no or virtually no scales).
 The N allele is lethal in the homozygous dominant state. The N allele has no effect on scale
pattern.
So far we have assumed that scaling pattern loci in carp only affect scale density and pattern.
However,allelic variation at a single locus may have effects on more than one biochemical
pathway and this can have subtle effects on other phenotypic characters. This is
called pleiotropy and there are a number of performance based pleiotropic effects on mirror,
line and leather carp. The wild-type scaled carp generally grow, survive and resist disease
better than do mirror, line or leather carp.
Figure. Inheritance of scale pattern in common carp.Scale pattern is determined by the
epistatic interaction between the S and N genes.
3.1.3.1 Dominant and recessive Epistasis
 Dominant epistasis occurs when a dominant allele at one locus (the epistatic locus)
produces a particular phenotype, regardless of the genotype at the second locus. The
second gene can express its phenotype only when the epistatic locus is homozygous recessive.
 For example, out of two genes, the dominant allele (e.g., A) of one gene masked the
activity of alleles of another gene (e.g., B) and expressed phenotypically, then A gene locus is
said to be epistatic to the B gene locus. Because, the dominant allele A can express itself only
in the presence of either B or b allele, therefore, such type of epistasis is termed as dominant
epistasis.
 The alleles of hypostatic locus or gene B will be able to express themselves
phenotypically only when gene locus Amay contain two recessive alleles (aa).

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 Thus, the genotype AABB or Aa Bb and AA bb or Aa bb produce the same
phenotype whereas the genotype aa BBor aa Bb and aa bb produce two additional
phenotypes. The dominant epistasis modify the classical ratio of 9:3:3:1 in to 12:3:1.
 In goldfish, two loci are involved simultaneously in the expression of
albinism. Albinism in goldfish is an example of a phenotype that is controlled by dominant
epistasis by the M and the S genes . M gene is the epistatic locus. A single dominant M allele
produces dark goldfish. When the M locus is homozygous recessive (mm), the S locus can
produce either light (SS, Ss) or albino (ss) goldfish.Consequently, albinos can only be
produced when a goldfish is homozygous recessive at both loci (mm, ss). and
 Recessive epistasis occurs when the recessive alleles of one gene locus (aa - the
epistasisl ocus) suppress the phenotypic expression of the alleles of another gene
(BB, Bb or bb alleles). This type of epistasis is calledrecessive epistasis. The alleles of B-
locus express themselves only when epistatic locus A has dominant allele
(e.g., AA or Aa). Recessive epistasis produces a 9:3:4 F2 phenotypic ratio. E.g. Eye colour
in Mexican cavecharacins. Black, brown and pink eye colour are controlled by
the ab and bw genes. ab locus is the epistatic locus–ab ab produces pink eyes, regardless of
the bw genotypes. A single dominant ab allele (+) allows the bwgenotype to produce either
brown or black eyes. F2 phenotype ratio for the mating of two heterozygous ab +, bw +black
eyed mexican cave characins are 9 Black : 3 Brown : 4 Pink. phenotype ratio.Table 5.2.
Phenotypes controlled by the different types of epistatic interaction of two genes in fish

Species Phenotype Type of epistasis F2 phenotypic ratio

Common carp Scale pattern Dominant epistasis 12:3:1
Duplicate recessive
Chinook salmon Flesh colour 9:7
gene interaction
Gold Fish Albinism Dominant epistasis 12:3:1
Mexican cave
Eye colour Recessive epistasis 9:3:4
characins
Duplicate genes with
Sumatran tiger barb Trunk striping 9:6:1
cumulative effects

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 3.1.4 Pleiotropism
Many and perhaps most of the biochemical pathways in the living organism are inter
connected and often inter dependent. Products of one reaction may be used in several other
metabolic schemes. Therefore, the phenotypic expression of a gene usually involves more
than one trait.
 The phenomenon of multiple effect (multiple phenotypic expression) of a single
gene is called pleiotropism. One gene has got its own effect on different parts or different
characteristics of one and the same organism.
 The characters that are governed by the single gene are not related. To a hatchery
manager, pleiotropic effects may be minor and insignificant if no economically important
characters are altered.
 However, when pleiotropic effects either increase or decrease viability, productivity
or market value, they become significant and can actually become more important than
phenotype. Pleiotropism has been extensively studied in common carp, Tilapia aurea, channel
catfish, etc.
3.1.4.1 Examples for pleiotropism in fishes
1. L, D, B and G colour genes in common carp have many pleiotropic effects. For e.g., Blue (bb)
and gold (gg) common carp have lowered growth rate as a pleiotropic effect.
2. Scale patterns in common carp are important qualitative phenotypes when they are grown for
food. The phenotypic effect of the S and N genes have been extensively studied. 17
pleiotropic effects were detected in mirror, line and leather carp. Some pleiotropic effects
include, mean growth rate of the fish, mean number of soft rays in dorsal fin and in anal fin,
mean number of rays in pelvic fin, erythrocyte count, etc.
3. The pleiotropic effect of gene N in common carp. One of the allelic pairs Nn (gene N) which
was lethal in the homozygous state reduced the growth rate, causing a reduction in size of
many organs and diminished the viability of the heterozygotes. Pleiotropic effect of gene N in
linear carp with genotype Nn includes the reduction of hemoglobin content and erythrocyte
number. These carps were less resistant to heating and oxygen deficiency. Gene S has a
pleiotropic influence on a host of characters, in particular on the structure of swim bladder,
but pleiotropy in this case is much weaker. Heterozygous for gene S are distinguished by a
slightly elevated viability. The degree of expression of the scale genes is markedly dependent
upon the presence of other modified genes.

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4. The pleiotropic effect of gene L in Japanese carp. “Light-coloured” carp result from a
dominant mutation frequently among the Japanese decorative carp. Carp, homozygous with
respect to the mutant gene (LL) die at the stages of the larvae or fry, no living fishes with this
genotype are found among the fingerlings; the heterozygotes (LI) survive but posses a
lowered viability. The gene ‘L’ in a heterozygote state has a pronounced pleiotropic action,
the pectoral fins undergo elongation, the posterior chamber of the air bladder becomes shorter,
and the head dimensions increase. The light-coloured carp have larger intestine, the protein
content in the blood serum is decreased. Their growth rate is characteristically accelerated to
about 20% during the first year of life and their behaviour is quieter. The lighter pattern of the
pigmentation throughout the trunk is due to the stable contraction of melanophores.
5. Dominant mutation found in koi is associated with the particular light-yellow pattern (the
stripe on the back and the ornamental pattern on the head). Carp with such a pattern, both
homo and hetero ones (DD and Dd) are rarely viable. The gene D is pleiotropic first like the
gene L, the head size is increased, the posterior chamber of the air bladder is elongated, the
number of vertebrae is increased.
6. The a allele in channel catfish produces albinism in the homozygous state (aa). Pleiotropy
effects of the ‘aa’ genotype are that albinos spawn later, produce smaller egg mass; produce
poorer quality eggs, have a poor percentage of hatching; produce progeny that are less viable
and have a poor growth rate than normally pigmented channel catfish. Moreover, because of
their brighter colour albinos are subject to more predation than normally pigmented channel
catfish.
7. The S allele in T. aurea produces saddle back in the heterozygous state (S+). Pleiotropic
effects of the S+ genotype are vertebral anomalis in vertebrae 1, 2 and 3; abnormal pelvic,
pectoral or anal fin, abnormal caudal skeletons; lowered disease resistance and reduced
viability.

3.1.5 Dominance
Complete dominant gene action
 Complete dominant gene action occurs when the dominant allele is so strong that it
produces its phenotype, regardless of the genotype.
 Only a single dominant and heterozygous genotypes produce the dominant
phenotype; thus, the phenotypes produced by these genotypes are identical.

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 The recessive allele can produce the recessive phenotype only when a fish is
homozygous recessive. Consequently, with this mode of inheritance, there are three genotypes
but only two phenotypes as described by Mendel.
 An example of a phenotype controlled by a single autosomal gene with complete
dominance is albino and normally pigmented channel catfish (Ictalurus punctatus). Normal
pigmentation is the dominant trait and is produced by the ++and +a genotypes. Albinism is
the recessive trait and is produced by the aa genotype.
 Table 5.3 lists some qualitative phenotypes in important cultured food fishes and
ornamental fishes that are produced by single autosomal genes with complete dominance.
Table 5.3. Phenotypes controlled by singleautosomal genes with complete dominant gene
action.

Species Allele Dominant phenotype Recessive phenotype

Channel catfish + a Normal pigmentation Albino

B b Normal pigmentation Blue

Common carp G g Normal pigmentation Gold
Gr gr Normal pigmentation Grey

Grass carp A a Normal pigmentation Albino

G g Normal pigmentation gold Albino

Rainbow trout
B b Normal pigmentation Metallic blue

B b Orange – red Blue

Goldfish D d Normal – eyes Telescope eyes
+ +ne Normal scales Nacreous – like

Curved spine
Sn Sc Normal spine
Blond soft yellow
Guppy B b Greenish grey
body colour
G g Greenish grey
Gold

F f Normal Fused vertebrae

Medaka
W w Normal spine Curved spine

Platyfish St st Stippled Unstippled

BL Bl Normal pigmentation Blond

Nile tilapia
b b Normal pigmentation light coloured (pink )

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 For example, the gold colour variant in common carp (Cyprinus carpio), is controlled at a
single locus by two alleles, a wild-type allele (G) and a variant gold producing allele (g). The
gold phenotype is only expressed in the fish when the g allele is present in the homozygous
condition (i.e. the genotype is gg),while the other two genotypes (GG andGg) will always
have wild-type colouration. The GG is present either in the homozygous or heterozygous
condition and is described as dominant. Conversely, the g allele is described as recessive
because its phenotypes is notexpressed in the heterozygous condition.
 If the gold variant is of high market value, the breeder can produce them in large numbers in
the following method.
 By mating a gg male with a gg female to ensure gg offspring; however, in wild population,
where are likely to be only occasional individuals expressing the gold coloration. So, to begin
with, if we mate our novel gold-colored fish to a wild-type fish, usually no gold offspring
would appear in the F1generation because the gg allele being present in both the father and the
mother, no gg genotypes–and therefore no gold phenotypes–will be present among the
F1offspring.
 However, the F1 offspring will be heterozygotes with one copy of the G allele and if these are
mated together then this will produce gg in F2 generation.
 “Blue”colour in carps of domesticated varieties is inherited as a simple recessive trait. This
blue color results from the under development of guanine crystals associated with the
reduction of guanophores in the skin of the carp. This condition is known as alampia.
3.1.5.1 Incomplete dominant gene action
A second type of dominance occurs when the dominant allele is incompletely dominant, i.e.,
the dominant allele always produces its phenotype, but it is unable to completely suppress the
recessive allele in the heterozygous state. When this happens, a gene with incomplete
dominant gene action (it has an incompletely dominant allele and a recessive allele) has three
genotypes,and each genotype produces a unique phenotype.
 The heterozygous genotype produces a phenotype that resembles but is slightly different from
the dominant phenotype. Because of this, the dominant phenotype can be produced only when
a fish has two copies of the dominant allele (homozygous dominant). Since the recessive
allele is not completely suppressed by the dominant allele, the heterozygous genotype
produces a phenotype that resembles, but is not identical to, the dominant phenotype.

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 In Black Lace angel fish there is a blending of traits, therefore Dark and Silver (wild type) act
as incomplete dominants to each other. (Table 5.4).
 More commonly, dominance may be incomplete, such that heterozygotes will express a
different colour to those expressed in the two homozygote classes. An example of partial
dominance is found with the ‘gold’ phenotype in the tilapia species Oreochromis
mossambicus, where the wild-type allele (G) is only partially dominant over the gold-type
allele (g) – GG genotypes are black, Gg heterozygotes have a ‘bronze’ skin colour
and gg homozygotes express the normal gold coloration. There are three genotypes and three
phenotypes, a unique phenotype for each genotype.
 Another example is the V gene in fighting fish. The V gene determines the number of
guanophores, and that in turn affects body colour (Table 5.4).
3 figures
Figure 5.4. Inheritance of black, bronze, and gold body colours inMozambiquetilapia.
Table 5.4. Phenotypes controlled by single autosomal genes with incomplete dominant gene
action.

Dominant Heterozygous Recessive

Species Allele
phenotype phenotype phenotype
Common carp L l Death Light coloured normal pigmented
Saddle back
Tilapia aurea S + death normal
(abnormal dorsal fin)
Mozambique tilapia G g black bronze gold
Goldfish T T1 transparent scalescalico normal scales
Siamese fighting fishV v steel blue blue green
Sail fin molly M m melanistic Spotted unspotted
Guppy Pl Pl+ death fused vertebrae normal vertebrae
long finned (full long finned (almost full
Angel Fish Lf + normal fish
phenotype) phenotype)

3.1.5.2 Codominant alleles

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In some situations the two alleles at a locus are co-dominant,i.e., both contribute equally to the
phenotypic character of the heterozygote. This inheritance pattern is difficult to distinguish
from the partial or incomplete dominance of a single allele, but fortunately the consequences,
from the point of view of the breeder, are the same.
 Obviously, to produce 100% heterozygote phenotype offspring, matings must occur
between two homozygotes.
 Alleles which lack dominant and recessive relationships may be called intermediate
or codominant alleles. This means that each allele is capable of some degree of expression
when in the heterozygous condition.
 Hence the heterozygous genotype gives rise to phenotype distinctly from either of the
homozygous genotypes. Usually heterozygous phenotype resulting from codominance is
intermediate in character between those produced by the homozygous genotypes, hence the
erroneous concept of “blending”. The phenotypes may appear to be a “blend” in heterozygotes
but the alleles maintain individual identities and will segregate from each other in the
formation of gametes.
 Example: Blood group system in human beings. A is dominant to O, B is dominant to
O, A and B are codominant and O is recessive to A and B.
3.1.5.3 Additive gene action
When there are more than 2 alleles, they all combine together and express a phenotype. When
the mode of gene action isadditive, neither allele is dominant, and each allele always
produces its phenotype in a unidirectional step-wise manner. When this occurs, a gene with
additive gene action produces a unique phenotype for each genotype:
 Only one additive gene has been discovered in fish; it controls golden, palomino, and
normal body colours in rainbow trout, Oncorhynchus mykiss.
 The G gene in rainbow trout is an example of a gene with additive gene action.
The G gene produces the golden, palomino, and normally pigmented body colour types. The
mating between two heterozygotes will produce progeny with 1:2:1 genotypic and phenotypic
ratios.

Genotype Phenotype
G1G1 Golden (red)

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 G1G Palomino (dark yellow)
GG Normally pigmented

 Additive interactions occur when two or more loci affect the same character. When
phenotypes are controlled by additive gene action, there is no dominant or recessive allele.
 Their effects are additive. Both alleles contribute equally to the production of the
phenotypes, so the heterozygous genotype produces a phenotype that is intermediate between
those produced by the two homozygous genotypes.
 Three phenotypes of the sticklebacks are controlled by additive action of 2 pairs of
genes A and a, B and b.
Phenotypes Strong Intermediate Weak
Genotypes AABB AaBB AABb aaBBAaBb AAbb aaBb Aabb aabb

3.1.6 Lethal genes

A gene whose phenotypic effect is sufficiently drastic to kill the bearer is called lethal gene.
Death from different lethal genes may occur at any time from fertilization of the egg to
advanced age.
 Lethal genes may be dominant, incompletely dominant or recessive.
 The fully dominant lethal gene kills the carrier individual both in homozygous and
heterozygous conditions.
 Completely dominant lethal genes usually cause death of the zygote later in
embryonic development or even after birth or hatching. No individuals will attain the age of
reproduction.
 Lethal genes arises occasionally by mutation from normal allele. The individual with
a dominant allele die before they can produce the progeny. Therefore the mutant dominant
allele is removed from the population in the same generation in which it arose.
 The recessive lethal allele kills the carrier individual only in homozygous condition.
 Certain lethal genes produce certain disorder to the individuals but do not destroy the
individual completely. They handicap but do not destroy. Such genes are called semi lethal
genes or sub lethal genes or subvital genes.

3.1.6.1 Dominant lethal genes

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Dominant lethal gene in common carp.
 Lethal genes N (reduction of scales) and L (lighter pigmentation) kill the carriers in
the homozygous state.
  The S gene in Tilapia aurea in another example of a dominant lethal gene.

Genotype Phenotype
SS death
S+ saddleback (abnormal dorsal fin)
++ normal

The mating of two saddleback (S+) T. aurea produces the expected 1:2:1 genotypic and
phenotypic ratios, but because all homozygous dominant fish are aborted, the genotypic ratio
2S+:1 ++ and 2 saddleback : 1 normal phenotypic ratios are observed.
3.1.6.2 Recessive lethal genes
 In guppy, Poecilia reticulata, when two Y-chromosomes are combined in a male,
homozygotes with respect to genesma (maculatus), ar (Armatus) and pa (Pauper) turn out to
be non-viable.
 Mean while, the males with genotypes Yma Yar, Yma Ypa and Ypa Yar are viable
and in addition fertile.
 It appears that a special lethal gene is closely linked with each of the genes located in
Y chromosome;
 the lethal genes associated with different genes are non-allelic.
3.1.6.3 Semi-dominant lethal gene
 The locus N (Nigra) in platy is a result of dominant mutation and is widely used by
aquarium fish culturists. The mutation results in a markedly blackening of the tail part of the
body and of the caudal fin. The pigmentation is due to the accumulation of large
melanophores.
 The semi dominant gene Fu producing the black pigmentation of the whole body
appears to be an allele of the geneN.
 The homozygotes Fu are non-viable. In the presence of recessive allele n the
macromelanophores are completely absent.

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  In angel fish, Pterophyllum scalare, there are about 25 colour patterns which are the
most important ornamental traits in this species. Golden body colour is determined by a
recessive autosomal gene designated by a semi-lethal autosomal gene, BI.
3.1.6.4 Sex-linked lethal gene
The X and Y chromosome of medaka. Oryzias latipes contain one “pigment” locus R with
three alleles.
 Normal fishes living in nature contain R gene in the sex chromosomes X and Y.
 Genetic analysis has shown that YR YR males obtained in crossing are practically non-
viable and the lethal ratio 2:1 has been reported.
3.1.7 Multiple Alleles
A type of inheritance unknown to Mendel is that called multiple allelism .
Some genes occur in more than two allelic forms (in contrast to alternative forms). In such
cases the various allelic forms are collectively referred to as multiple alleles. A set of multiple
alleles may contain three, four and even upto twenty or more members and all the members
essentially occupy the same locus in homologous chromosomes.
 It therefore follows that regardless of their total number, only two members of a set
occur in a diploid cell, and only one in a gamete (haploid cell).
 All the members of a set of multiples alleles are infact the mutant forms of the same
gene. Since mutations keep occurring the membership of a set is subject to increase.
 In a set of multiple alleles, one member is always dominant to all the others and one
member will be always recessive to all the others.
 The presence of multiple alleles is also reported in fishes.
1. The B gene which controls melanin formation in the Medaka’s melanophores, is an
example of an autosomal gene with three alleles. The B allele is dominant over
the B’ and b alleles and the B’ allele is dominant over the b allele; the b allele is recessive to
the other two.

Genotype Phenotype
BB BB Bb Full melanin production
B’ B’ B’b Variegated pigmentation
bb No or minimal melanin pigmentation

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2. In platy fish there are nine alleles at the P locus, P+, PM, PMC, Pr, PCO, PC, PCc, PO, PD

P+ - unpotted phenotype which is recessive to all other alleles.

 The remaining alleles are co-dominant. They produce both phenotypes.
 The alleles at the P locus can in theory produce 37 phenotypes but a number of patterns
overlap which reduces the number of visible phenotypes to 27.
PM – Moon, PMC- -Moon complete, Pr – twin spot, PCO - comet, PC- crescent, PCc- complete
crescent- PO - one spot, PD - dot.
3.1.8 Complementary genes or duplicate recessive genes (9:7)
If both gene loci have homozygous recessive alleles and both of them produce identical
phenotypes, theF2 ratio 9:3:3:1 would become 9:7. Both dominant alleles when present
together (AA BB, Aa BB, AA Bb, Aa Bb) and complement to each other and are called
complementary genes and produce different phenotype.
 This 9:7 ratio is observed in ‘Pearl’ Nile tilapia.
 The ‘pearl’ character of Nile tilapia (O. niloticus) where the phenotypic opalescent-
white character of the scales (designated ‘pearl’) is also controlled epistatically at two loci.
 Two loci A and B are involved. Each locus has two alleles. Normal colouration is
produced by A and B. Alleles a and b are recessive.
 The a and b alleles in combination, and only in combination, would result in pearl
colouration. Either recessive allele, when present alone, would not alter the normal

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colouration of the fish, but if one copy of each a and b were present, the result would be pearl
colouration.
 This is illustrated in a Punnett square where the wild-type alleles at the two loci
are A and B and the pearl-type alleles are designated a and b respectively.

Gametes AB Ab aB ab

AA BB AA Bb Aa BB Aa Bb
AB
wild-type wild-type wild-type pearl
AA Bb AA bb Aa Bb Aa bb
Ab
wild-type wild-type pearl pearl
Aa BB Aa Bb Aa BB aa Bb
aB
wild-type pearl wild-type pearl
Aa Bb Aa bb Aa Bb aa bb
ab
pearl pearl pearl pearl

Ratio of phenotypes: 7wild-type : 9 pearl

Figure 5.5. Punnett square illustrating inheritance of the ‘pearl’ phenotype in the Niletilapia
(O. niloticus) based on variation at two epistatic loci. Matings were between doubly
heterozygous F1 individuals (Aa Bb x Aa Bb).