GENETICS LECTURE NOTE

ND TWO SLT AND BIOTECH

BY

Dr. Abdulhadi Yakubu

(Ph.D Industrial Microbiology)

BASIC CONCEPTS OF GENETICS

All living organisms reproduce which leads to the formation of offspring of the same kind.
Example, a pea plant produces only pea plants each time it reproduces, rat produces only rats and
humans produce only humans. However, the resulting offspring need not and most often do not
totally resemble the parent. Several characteristic differences may occur between individuals
belonging to the same species. The similarities and differences among the members of a species
are not coincidental. Both the similarities and differences have been received from their parents.
The mechanism of transmission of characters, resemblances as well as differences, from the
parental generation to the offspring, is called as HEREDITY. The differences shown by
individuals within the same species and in the offspring are described as VARIATIONS. The
scientific study of heredity, variations and the environmental factors responsible for these, is
known as GENETICS.
Genetics is simply the study of the function and behavior of genes. Genes are bits of biochemical
instructions found inside the cells of every organism from bacteria to humans. Offspring receive
a mixture of genetic information from both parents. This process contributes to the great

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variation of traits that we see in nature, such as the color of a flower’s petals, the markings on a
butterfly’s wings, or such human behavioral traits such as personality or musical talent.
Geneticists seek to understand how the information encoded in genes is used and controlled by
cells and how it is transmitted from one generation to the next.
Genetic information is encoded and transmitted from generation to generation in
deoxyribonucleic acid (DNA). DNA is a coiled molecule organized into structures called
chromosomes within cells. Segments along the length of a DNA molecule form genes. Genes
direct the synthesis of proteins, the molecular laborers that carry out all life-supporting activities
in the cell. Although all humans share the same set of genes, individuals can inherit different
forms of a given gene, making each person genetically unique

APPLICATION OF GENETICS TO SOCIETY

• Genetics is central to the life of every individual: it influences our physical features,
susceptibility to numerous diseases, personality, and intelligence.
• Genetics plays important roles in agriculture, the pharmaceutical industry, and medicine.
 It is central to the study of biology.
• Genetic variation is the foundation of evolution and is critical to understanding all life.
• The use of genetics by humans began with the domestication of plants and animals

GENES AND THEIR APPLICATIONS

1. A gene is the fundamental unit of heredity- The precise way in which a gene is defined
often varies. At the simplest level, we can think of a gene as a unit of information that
encodes a genetic characteristic.
2. Genes come in multiple forms called alleles- A gene that specifies a characteristic may
exist in several forms, called alleles. For example, a gene for coat color in cats may exist
in alleles that encode either black or orange fur.
3. Genes encode phenotypes- One of the most important concepts in genetics is the
distinction between traits and genes. Traits are not inherited directly. Rather, genes are
inherited and, along with environmental factors, determine the expression of traits. The
genetic information that an individual organism possesses is its genotype. The trait is its

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 phenotype. For example, the A blood type is a phenotype; the genetic information that
encodes the blood type A antigen is the genotype.
4. Genetic information is carried in DNA and RNA: Genetic information is encoded in
the molecular structure of nucleic acids, which come in two types: Deoxyribonucleic
Acid (DNA) and Ribonucleic acid (RNA). Nucleic acids are polymers consisting of
repeating units called nucleotides; each nucleotide consists of a sugar, a phosphate, and a
nitrogenous base. The nitrogenous bases in DNA are of four types which includes
adenine, cytocine, guanine and thiamine (abbreviated as A, C, G, and T), and the
sequence of these bases encodes genetic information. Most organisms carry their genetic
information in DNA, but a few viruses carry it in RNA. The four nitrogenous bases of
RNA are adenine, cytocine, guanine and uracil (abbreviated A, C, G, and U).
5. Genes are located on chromosomes- The vehicles of genetic information within the cell
are chromosomes, which consist of DNA and associated proteins. The cells of each
species have a characteristic number of chromosomes; for example, bacterial cells
normally possess a single chromosome; human 46 and pigeon 80.
CHROMOSOMES
Chromosomes (chroma = colour, soma = body) are tiny thread-like structures found in the
nucleus of a cell. In a non-dividing cell, they appear as a chromatin network while during cell
division they become condensed to form short and thick chromosomes. Chromosomes are unique
cell structures which are capable of replication. They store and transmit the coded information
which is responsible for all the life processes of an organism. Hence, chromosomes are
commonly described as carriers of heredity. The term "chromosome" was coined by Waldeyer in
1888 since these structures easily take up dye stains.
THREE ESSENTIAL ELEMENTS OF CHROMOSOME

1. CENTROMERE:

This is the attachment point for spindle microtubules, which are the filaments responsible for
moving chromosomes during cell division. The centromere appears as a constricted region that
often stains less strongly than does the rest of the chromosome. Before cell division, a protein
complex called the kinetochore assembles on the centromere, to which spindle microtubules later
attach.

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2. A PAIR OF TELOMERES:

• Are the natural ends, the tips, of a linear chromosome.

• They serve to stabilize the chromosome ends. If a chromosome breaks, producing new
ends, these ends have a tendency to stick together, and the chromosome is degraded at the
newly broken ends.

• Telomeres provide chromosome stability.

• Some research suggest that telomeres also participate in limiting cell division

• Play important roles in aging and cancer

3. ORIGINS OF REPLICATION

• Are the sites where DNA synthesis begins

• They are not easily observed by microscopy.

Types of chromosomes
a) based on types of organism

1. Bacterial chromosome (prokaryotic)

In bacteria, which being prokaryotic organisms, the entire hereditary material are packed
into a single, irregularly folded compact mass called nucleoid or genophore or bacterial
chromosome. It is short and simple consisting of a single DNA molecule. The DNA is in
the form of a double helix which forms a closed ring or circle with no free ends. It is
permanently attached to a mesosome, an infolding of the plasma membrane. The bacterial
chromosome lacks a protein coat and it is in direct contact with the cytoplasm, since a
nuclear membrane is absent. A small amount of protein, mainly in the form of an enzyme
called RNA polymerase, may be found associated with the bacterial chromosome.
2. Eukaryotic chromosome

The eukaryotic cells show a varied number of chromosomes. It is presumed to be the result
of breaking up of a single long chromosome into several short units, in the course of

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evolution, to accommodate the increase in the amount of genetic information. The eukaryotic
chromosomes occur inside the nucleus of a cell, separated from the cytoplasm by a distinct
nuclear membrane.

b) Based on autosome and sex chromosome

1. Autosomes (somatic)

• Carry genes which determine the somatic characteristics

• Do not have any influence on determining the sex of the organism

2 Allosome (Sex chromosome)

 Carry genes responsible for sexual characteristics
 Have a significant role in the determination of sex.

c) On the Basis of Number of Centromeres

• Monocentric (single centromere) found in many plants and animals

• Dicentric (Two centromeres) found in plants such as maize

• Polycentric (many centromeres) found in Roundworms. E.g. Parascaris equorum

• Holocentric (acentric): Lack centromere. The entire surface of the chromatid will
fuction as centromere. E.g. in nematode like Caenorhabditis elegans

d) On the Basis of Location of Centromere

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Function and Significance of Chromosomes

Genetic Code Storage: Chromosome contains the genetic material that is required by the
organism to develop and grow. DNA molecules are made of chain of units called genes.
Genes are those sections of the DNA which code for specific proteins required by the cell for
its proper functioning.

Sex Determination: Humans have 23 pairs of chromosomes out of which one pair is the sex
chromosome. Females have two X chromosomes and males have one X and one Y
chromosome. The sex of the child is determined by the chromosome passed down by the
male. If X chromosome is passed out of XY chromosome, the child will be a female and if a
Y chromosome is passed, a male child develops.

Control of Cell Division: Chromosomes check successful division of cells during the process
of mitosis. The chromosomes of the parent cells insure that the correct information is passed
on to the daughter cells required by the cell to grow and develop correctly.

Formation of Proteins and Storage: The chromosomes direct the sequences of proteins
formed in our body and also maintain the order of DNA. The proteins are also stored in the
coiled structure of the chromosomes. These proteins bound to the DNA help in proper
packaging of the DNA.

GENETIC TERMS AND MENDEL’S EXPERIMENTS

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1. Phenotype- this is the physical expression of genes in an organism. It is denoted with
words such as Tall, Dwarf or Short, Smooth, Wrinkled, Long wing, Vestigial wing, etc.
2. Genotype- The genetic constitution of a cell or an organism, as distinguished from its
physical and behavioral characteristics, i.e., its phenotype. It is denoted with letters such
as TT or Tt for Tall, SS or Ss for Smooth, etc. The genotype of an organism may be
homozygous or heterozygous.
3. Allele or Allelomorphic pair- A pair of contrasting genes controlling a character. The
pair could be identical- T&T or different-T&t
4. Homozygous-individual with identical alleles (TT , tt, SS, ss) controlling same character
or trait. Such organisms always produce identical gametes during meiosis and are thus
said to be pure breeding or true breeding or breeds true.
5. Heterozygous- Individual with non-identical or dissimilar alleles (Tt, Ss,Rr) controlling
same character or trait. Such organisms produce different or non-identical gametes during
meiosis and are thus said not to be pure breeding or true breeding or does not breed true.
6. Dominant trait or character -this is controlled by dominant genes that can express itself
in all generations in the presence of the contrasting gene. It suppresses the effect of the
contrasting gene. It is denoted with capital letters such as TT , SS, etc
7. Recessive trait or character -this is controlled by recessive genes that can not express
itself in all generations, but only in certain generations in the absence of the dominant
gene. It is denoted with small letters such as tt, ss.
8. Filial generations - these are the series of offspring produced from genetic crossings. It
is denoted with letter F and a subscript to show the particular generation, e g F1, F2 , F3
for first , second and third Filial generations respectively.
9. Hybrid - a product of crossing between two contrasting parents. For example, Tall x
Short to produce All Tall F1 offspring. That is TT x tt to give Tt.
10. Monohybrid cross- A cross between a pair of contrasting characters, e.g. Tall x Short,
Smooth pod x Wrinkled pod, etc. At F1 the dominant character is expressed in all the
offspring while the recessive character is masked at F2
11. , the genotypic ratio of the offspring is 1: 2:1 while the phenotypic ratio is 3:1 in
complete dominance condition.

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 12. Dihybrid cross- a cross between two pairs of contrasting characters, e.g. Tall &Smooth x
Short and Wrinkled. At F1, the dominant character is expressed in all the offspring, while
the recessive character is masked. However, at F2, both the dominant and recessive
characters are expressed in the offspring in varying genotypic ratios and phenotypic ratio
of 9:3:3:1.

Brief history of genetics

The biggest name in the entire history of genetics beyond any doubt is that of Gregor Johann
Mendel (1822-1884). It was Mendel, more than any other scientist, who synthesized the basic
principles of heredity into a body of knowledge that has formed the very core of modern
genetics. It must be emphasized here that Mendel was not the original pioneer in the field of
genetics. As with all other scientific achievements, many scientists before the period of Mendel
had laid the foundation. However, it was Mendel who combined the ideas put forth by other
scientists into a definite set of working principles that are acceptable even today. Like so many
other discoveries by famous scientists, Mendel's ideas on the mechanism of inheritance did not
gain any importance and his principle temporality died with him. Fortunately for science, three
scientists, Hugo De Vries, Tschermarck and Correns rediscovered the ideas, when they obtained
the same results in the experiments conducted by each one of them independently. The
rediscovery of Mendelism brought new emphasis to the field of heredity and the modern science
of genetics was born. The basic ideas and the conclusions drawn by scientists after this
rediscovery came to be known as Mendelian Genetics. It was during this period that Mendel
developed curiosity over the pattern of inheritance of characters from parent organisms to
offspring. After careful thought, he designed breeding experiments in the pea plants. He carefully
analyzed the results, gave a mathematical interpretation and published them in 1866. However,
unfortunately for Mendel, his results and conclusions could not convince the contemporary
biologists. Mendel died in 1884 without knowing that he had laid the foundation for modern
genetics. It was only in 1900, that Mendel's work was rediscovered and its significance was
made known to the scientific world. Nevertheless, Mendel is now regarded as the father of
modern genetics' for his significant and pioneering contributions to the field of genetics.

S/No. Parameters Contrasting characters

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 1 Length of the stem
2 Position of the flowers Tall and short
3 Nature of the fruit Axial and terminal
4 Color of the unripe fruit Inflated and constricted
5 Color of the seed coat Grey and white
6 Nature of the seed coat Round and wrinkled
7 Color of the cotyledons Yellow and green

Monohybrid Inheritance
In the initial set of experiments, Mendel concentrated only on the pattern of inheritance of a
single pair of contrasting characters. This pattern of inheritance involving only one pair of
contrasting characters is known as monohybrid inheritance. In the first set of experiments,
Mendel conducted cross-pollination between a pure breeding tall plant and a pure breeding dwarf
plant. He collected the seeds from this cross pollination and allowed them to germinate. All the
resulting plants were found to be tall.

Based on these results, Mendel came to the conclusion that in a cross-involving two contrasting
characters, only one character expresses itself in the next generation. Mendel called the
character, which expressed as dominant character and the character, which failed to express, as

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recessive character. At this stage, Mendel wanted to know whether the tall plants resulting from
a cross between tall and dwarf plants, were similar to the tall plants of the P 1 generation. Hence,
he allowed the tall plants of the F1 generation to undergo self-pollination. In the next generation,
Mendel found both tall plants and dwarf plants, approximately in the ratio 3:1. The results were
most surprising since the recessive character dwarfness had reappeared in the next generation (F 2
generation). From the results, it was clear that the tall plants of the F1 generation were different
genetically from the tall plants of the P1 generation. Similar results were obtained by Mendel for
the other contrasting characters also. Based on these results, Mendel came to the conclusion that
certain factors are involved in the expression of each of these contrasting characters. He
presumed that if the F1 tall plants on self pollination could give rise to both tall and dwarf plants,
this plant should have contained two factors, one responsible for tallness and the other
responsible for dwarfness.
Similarly, he presumed that the P1 tall plants also should have contained two factors, both
responsible for tallness. He represented these ideas by using the letters of English alphabet to
represent the factors. He represented the factor for dominant character by a capital letter and the
factor for recessive character by a small letter.
Eg : Factor for tallness ------------T
Factor for dwarfness -------- t
PUNNET SQUARE
The genotypes and phenotypes resulting from various combinations of gametes can be easily
determined by Punnet squares, devised by Reginald C. Punnet (1875 1967). Hence each of the
possible gametes is placed in an individual column or a row, with vertical column representing
the female and horizontal row the male parent. The gametes are then arranged in all possible
combinations and the resulting genotypes are entered in the boxes along with the phenotypes.

Tall plant Dwarf plant

TT tt
Gametes T T x t t

T t
T Tt Tt

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 T Tt Tt
All tall plants

MENDELIAN LAWS OF INHERITANCE

Mendel’s First Law of inheritance:
This first law is also called the law of segregation of genes. The law states that, genes are
responsible for the development of the individual and that, they are independently transmitted
from one generation to another without undergoing any alteration.
Mendel’s second Law of Inheritance
This second law is also called the law of independent assortment. This law states that gametes
produced by segregation of genes as explained by Mendel’s first law behaves as a separate unit
and are inherited independently of any other gametes. That is, each of the gametes has equal
chances of mating with one another.
Test Cross and Back Cross
Mendel devised a system of conducting verification for the results obtained by him. It is known
as test cross. It is a cross between F1 plant and the recessive parent. A test cross conducted for
the monohybrid inheritance results in the two opposite characters expressing in a ratio of 1:1.
Significance of Test Cross
1. Test cross can be used to determine the genotype of the F1 plant.
2. The test cross can be used to support the idea that the reappearance of the recessive character
in the F2 generation is due to the heterozygous condition of the F1 plant.
3. The test can be used to verify whether any given pair of characters can be alleles (contrasting
characters).
BACK CROSS
If an F1 individual or an individual of F2 or F3 generations is crossed with any one of the parents
it is called a back cross
Monohybrid inheritances in fruit fly (Drosophila melanogaster)

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The fruit fly Drosophila melanogaster, has eight chromosomes: three pairs of autosomes and one
pair of sex chromosomes. The fruit fly can easily be cultured at 25°C. It completes its
development in two weeks. This means that several generations can be studied in a short time.
The wild fly can he crossed to various mutant forms. E.g. flies with vestigial wings.
Let the alleles be represented by symbols
VG = dominant gene for long wing
vg = recessive gene for vestigial wing

In the above monohybrid experiment, two alleles for wing length are involved. The female
parent is homozygous for the gene responsible for the long wing while the male parent is
homozygous for the recessive character and exhibits vestigial wings as its phenotype. During
gamete formation, the genes separate and each gamete carries only one of the two genes. After
fertilization the resulting zygote possesses one gene of each type and they are therefore termed
heterozygous. Since the gene for long wing is dominant they all manifest this character.
DIHYBRID INHERITANCE
Mendel did not limit his experiments to testing the rules of inheritance of single traits. He also
studied plant traits involving multiple pairs of genes, breeding plants that have round, yellow
seeds with plants that produce wrinkled, green seeds. Such experiments demonstrated that the
patterns of inheritance he observed in his experiments with single traits also apply to cases
involving more complex gene combinations. In addition to his work on monohybrid crosses,
Mendel also crossed varieties of peas that differed in two characteristics (dihybrid crosses). For
example, he had one homozygous variety of pea plant that produced round seeds and yellow

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endosperm; another homozygous variety produced wrinkled seeds and green endosperm. When
he crossed the two, all the F1 progeny had round seeds and yellow endosperm.
He then self-fertilized the F1 and obtained the following progeny in the F2: 315 round, yellow
seeds; Mendel recognized that these traits appeared approximately in a 9:3:3:1 ratio; that is, of
the progeny were round and yellow, were wrinkled and yellow, were round and green, and were
wrinkled and green.
Lets YY be yellow color and yy be green color
RR be round seed and rr be wrinkled seed
Therefore, round yellow seed is RRYY
While wrinkled green seed is rryy
Parents are RRYY and rryy
Gametes RY, RY and ry, ry

X RY RY
Ry RYry RYry
Ry RYry RYry

At F1, all the fruit are physically round yellow but genetically heterozygous round yellow

When a cross involving F1 here, F2 will produce 16 different fruit as follows

RYry cross with RYry

Gamete from each fruit is RY, rY, Ry, ry and RY, rY, Ry, ry
We also use punnet square for the cross as below
X RY rY Ry Ry
RY RRYY RrYY RRYy RrYy
rY RrYY rrYY RrYy rrYy
Ry RRYy RrYy RRyy Rryy
Ry RrYy rrYy Rryy Rryy

Phenotypic ratio is 9:3:3:1

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Genotipic ratio includes
1. Homozygous round homozygous yellow (RRYY)………….1
2. Heterozygous round homozygous yellow (RrYY)…………..2
3. Homozygous round heterozygous yellow (RRYy)…………….2
4. Heterozygous round heteroxygous yellow (RrYy)……………..4
5. Homozygous wrinkled homozygous yellow (rrYY)……………1
6. Homozygous wrinkled heteroxygous yellow (rrYy)……………2
7. Homoxygous round homoxygous green (RRyy)………………..1
8. Hereroxygous round homozygous green (Rryy)…………………2
9. Homozygous wrinkled homoxygous green (rryy)………………..1
Deviations from Mendelian ratio
(a) Incomplete Dominance
In cases of incomplete dominance, the inheritance of a dominant and a recessive allele results in
a blending of traits to produce intermediate characteristics. For example, four-o’clock paint
plants may have red, white, or pink flowers. Plants with red flowers have two copies of the
dominant allele R for red flower color (RR). Plants with white flowers have two copies of the
recessive allele r for white flower color (rr). Pink flowers result in plants with one copy of each
allele (Rr), with each allele contributing to a blending of colors.
(b) Quantitative Inheritance
Mendel focused his studies on traits determined by a single pair of genes, and the resulting
phenotype was easy to distinguish. A tall plant can be markedly different from a short one, and a
green pea can easily be distinguished from a yellow one. There are some traits, however, that are
not easy to distinguish. Human skin color, for example, may be any of a wide variety of shades.
Traits such as skin color differ from the ones Mendel studied because they are determined by
more than one pair of genes. In this form of inheritance, known as quantitative inheritance, each
pair of genes has only a slight effect on the trait, while the cumulative effect of all the genes
determines the physical characteristics of the trait. At least four pairs of genes control human
skin color. Multiple genes also control many traits important in agriculture, such as milk
production in cows and ear length in corn.
(c) Multiple Alleles

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Another exception to Mendelian genetics involves genes with multiple alleles. Certain traits are
controlled by multiple alleles that have complex rules of dominance. In humans, for example, the
gene for blood type has three alleles: IA, IB, and i. With three alternatives for each member of a
gene pair, there are six possible combinations of these genes (IAIA, IBIB, ii, IAi, IBi, IAIB). Although
there are six possible combinations, humans have only four major blood types: A, B, AB, and O.
This results because both IA and IB dominate over i, but not over each other, so a person with a
gene combination of IAIA or IAi has blood type A. The gene combinations IBIB and IBi both
produce blood type B. IAIB results in a blood type AB, and ii results in blood type O.
Blood Transfusion
The transfer of blood from one person to another is called blood transfusion. In all cases of blood
transfusion, it is necessary to match the blood group of the recipient with the blood group of
donor. The following table represents the blood group matching.

Donor Recipient A B (anti-a) AB (Nil) O (anti-a and

(anti-b) anti-b)
A (Antigen A) Yes No Yes No
B (Antigen B) No Yes Yes No
AB (Both A and B) No No Yes No
O (Nil) Yes Yes Yes Yes

From the table it is clear that persons with blood group AB can receive blood from any other
person. Hence they are commonly described as universal recipients persons with blood group O
can donate blood to any other person. Hence, they are commonly described as universal donors.

Blood groups and their genotypes

Blood groups Genotypes
A IAIA or IAi
B IBIB or IBi
AB IAIB
O Ii

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Sex Determination
Sexual reproduction is the formation of offspring that are genetically distinct from their parents;
most often, two parents contribute genes to their offspring. Among most eukaryotes, sexual
reproduction consists of two processes that lead to an alternation of haploid and diploid cells:
meiosis produces haploid gametes, and fertilization produces diploid zygotes. The term sex
refers to sexual phenotype. Most organisms have only two sexual phenotypes: male and female.
There are many ways in which sex differences arise. In some species, both sexes are present in
the same individual, a condition termed hermaphroditism; organisms that bear both male and
female reproductive structures are said to be monoecious (meaning “one house”). Species in
which an individual has either male or female reproductive structures are said to be dioecious
(meaning “two houses”). Humans are dioecious. Among dioecious species, the sex of an
individual may be determined chromosomally, genetically, or environmentally.
1. Chromosomal Sex-Determining Systems

a. XX-XO sex determination

The mechanism of sex determination in the grasshoppers studied by McClung is one of the
simplest mechanisms of chromosomal sex determination and is called the XX-XO system. In this
system, females have two X chromosomes (XX), and males possess a single X chromosome
(XO). There is no O chromosome; the letter O signifies the absence of a sex chromosome. In
meiosis in females, the two X chromosomes pair and then separate, with one X chromosome
entering each haploid egg. In males, the single X chromosome segregates in meiosis to half the
sperm cells—the other half receive no sex chromosome. Because males produce two different
types of gametes with respect to the sex chromosomes, they are said to be the heterogametic sex.
Females, which produce gametes that are all the same with respect to the sex chromosomes, are
the homogametic sex. In the XX-XO system, the sex of an individual is therefore determined by
which type of male gamete fertilizes the egg. X-bearing sperm unite with X-bearing eggs to
produce XX zygotes, which eventually develop as females. Sperm lacking an X chromosome
unite with X-bearing eggs to produce XO zygotes, which develop into males.
b. XX-XY sex determination

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In many species, the cells of males and females have the same number of chromosomes, but the
cells of females have two X chromosomes (XX) and the cells of males have a single X
chromosome and a smaller sex chromosome called the Y chromosome (XY). In humans and
many other organisms, the Y chromosome is acrocentric not Y shaped as is commonly assumed.
In this type of sex-determining system, the male is the heterogametic sex—half of his gametes
have an X chromosome and half have a Y chromosome. The female is the homogametic sex—all
her egg cells contain a single X chromosome. Many organisms, including some plants, insects,
and reptiles, and all mammals, have the XX-XY sex-determining system. Although the X and Y
chromosomes are not generally homologous, they do pair and segregate into different cells in
meiosis.
c. ZZ-ZW sex determination
In this system, the female is heterogametic and the male is homogametic. To prevent confusion
with the XX-XY system, the sex chromosomes in this system are labeled Z and W, but the
chromosomes do not resemble Zs and Ws. Females in this system are ZW; after meiosis, half of
the eggs have a Z chromosome and the other half have a W. Males are ZZ; all sperm contain a
single Z chromosome. The ZZ-ZW system is found in birds, moths, some amphibians, and some
fishes. Receiving the same allele from their mother and a 100% chance of receiving the same
allele from their father; the average relatedness between sisters is therefore 75%. Brothers have a
50% chance of receiving the same copy of each of their mother’s two alleles at any particular
locus; so their average relatedness is only 50%.
d. Haplodiploidy
Some insects in the order Hymenoptera (bees, wasps, and ants) have no sex chromosomes;
instead, sex is based on the number of chromosome sets found in the nucleus of each cell. Males
develop from unfertilized eggs, and females develop from fertilized eggs. The cells of male
hymenopterans possess only a single set of chromosomes (they are haploid) inherited from the
mother. In contrast, the cells of females possess two sets of chromosomes (they are diploid), one
set inherited from the mother and the other set from the father. The haplodiploid method of sex
determination produces some odd genetic relationships. When both parents are diploid, siblings
on average have half their genes in common because they have a 50% chance of receiving the
same allele from each parent. In these insects, males produce sperm by mitosis (they are already
haploid); so all offspring receive the same set of paternal genes.

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 2. Genic Sex-Determining Systems
In some plants and protozoan, sex is genetically determined, but there are no obvious differences
in the chromosomes of males and females (there are no sex chromosomes). These organisms
have genic sex determination; genotypes at one or more loci determine the sex of an individual.
It is important to understand that, even in chromosomal sex-determining systems, sex is actually
determined by individual genes. For example, in mammals, a gene located on the Y chromosome
determines the male phenotype. In both genetic sex determination and chromosomal sex
determination, sex is controlled by individual genes; the difference is that, with chromosomal sex
determination, the chromosomes that carry those genes appear different in males and females.
3 Environmental Sex Determinations
In this system, sex is determined fully or in part by environmental factors in a number of
organisms. One fascinating example of environmental sex determination is seen in the marine
mollusk Crepidula fornicata, also known as the common slipper limpet. Slipper limpets live in
stacks, one on top of another. Each limpet begins life as a swimming larva. The first larva to
settle on a solid, unoccupied substrate develops into a female limpet. It then produces chemicals
that attract other larvae, which settle on top of it. These larvae develop into males, which then
serve as mates for the limpet below. After a period of time, the males on top develop into
females and, in turn, attract additional larvae that settle on top of the stack, develop into males,
and serve as mates for the limpets under them. Limpets can form stacks of a dozen or more
animals; the uppermost animals are always male. This type of sexual development is called
sequential hermaphroditism; each individual animal can be both male and female, although not at
the same time. Environmental factors are also important in determining sex in many reptiles.
Although most snakes and lizards have sex chromosomes, in many turtles, crocodiles, and
alligators, temperature during embryonic development determines sexual phenotype. In turtles,
for example, warm temperatures produce females during certain times of the year, whereas cool
temperatures produce males. In alligators, the reverse is true.
The role of sex chromosomes
1. The X chromosome contains genetic information essential for both sexes; at least one copy of
an X chromosome is required for human development.
2. The male-determining gene is located on the Y chromosome. A single copy of this
chromosome, even in the presence of several X chromosomes, produces a male phenotype.

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3. The absence of the Y chromosome results in a female phenotype.
4. Genes affecting fertility are located on the X and Y chromosomes. A female usually needs at
least two copies of the X chromosome to be fertile.
5. Additional copies of the X chromosome may upset normal development in both males and
females, producing physical and mental problems that increase as the number of extra X
chromosomes increases.
Sex-Linked Characteristics
Sex-linked characteristics are determined by genes located on the sex chromosomes. Genes on
the X chromosome determine X-linked characteristics; those on the Y chromosome determine Y
linked characteristics. Because little genetic information exists on the Y chromosome in many
organisms, most sex-linked characteristics are X linked. Males and females differ in their sex
chromosomes; so the pattern of inheritance for sex-linked characteristics differs from that
exhibited by genes located on autosomal chromosomes Sex-linked linked inheritance can be seen
in eye colour in Drosophila melanogaster , colour blindness and haemophilia in man:
1. X-Linked Color Blindness in Humans
Within the human eye, color is perceived in light-sensing cone cells that line the retina. Each
cone cell contains one of three pigments capable of absorbing light of a particular wavelength;
one absorbs blue light, a second absorbs red light, and a third absorbs green light. The human eye
actually detects only three colors (red, green, and blue) but the brain mixes the signals from
different cone cells to create the wide spectrum of colors that we perceive. Each of the three
pigments is encoded by a separate locus; the locus for the blue pigment is found on chromosome
7, and those for green and red pigments lie close together on the X chromosome. The most
common types of human color blindness are caused by defects of the red and green pigments; we
will refer to these conditions as red–green color blindness. Mutations that produce defective
color vision are generally recessive and, because the genes coding for the red and green pigment
are located on the X chromosome, red–green color blindness is inherited as an X-linked
recessive characteristic.
We will use the symbol Xc to represent an allele for red–green color blindness and the symbol X
to represent an allele for normal color vision. Females possess two X chromosomes; so there are
three possible genotypes among females: XX, XcX and which produce normal vision, and XcXc,

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which produces color blindness. Males have only a single X chromosome and two possible
genotypes: XY, which produces normal vision, and XcY which produces color blindness.
If a color-blind man mates with a woman homozygous for normal color vision, all of the gametes
produced by the woman will contain an allele for normal color vision. Half of the man’s gametes
will receive the X chromosome with the color-blind allele, and the other half will receive the Y
chromosome, which carries no alleles affecting color vision. When an Xc-bearing sperm unites
with the X bearing egg, a heterozygous female with normal vision (XXc) is produced. When a
Y-bearing sperm unites with the X bearing egg, a heterozygous male with normal vision
(XY) is produced.
In the reciprocal cross between a color-blind woman and a man with normal color vision the
woman produces only Xc bearing gametes. The man produces some gametes that contain the X
chromosome and others that contain the Y chromosome. Males inherit the X chromosome. From
their mothers; because both of the mother’s X chromosomes bear the Xc allele in this case, all
the male offspring will be color blind. In contrast, females inherit an X chromosome from both
parents; thus the female offspring of this reciprocal cross will all be heterozygous with normal
vision. Females are color blind only when color-blind alleles have been inherited from both
parents, whereas a color-blind male need inherit a colorblind allele from his mother only; for this
reason, color blindness and most other rare X-linked recessive characteristics are more common
in males. In these crosses for color blindness, notice that an affected woman passes the X-linked
recessive trait to her sons but not to her daughters, whereas an affected man passes the trait to his
grandsons through his daughters but never to his sons.
Haemophilia (bleeding disease)
Haemopilia is an abnormality controlled by a recessive gene located on the X chromosome.
Bleeding from a puncture or an open wound takes an abnormally long time to stop or fails to stop
because clotting of blood would not occur. Small injuries like puncture, extraction of tooth and
the like can cause such persons to bleed to death.
Relevance of Genetics in disputed paternity
1) Suppose a child is of blood group AB and the mother of blood group A, Can a man of blood
group O be the father of such a child?
Solution:
The child of blood group AB will have the genotype IAIB

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The mother of blood group A will be of genotype IAIA or IAi
A man of blood group O will have the genotype ii
The possible crossings are:
Phenotypes O x A
Genotypes ii x IAIA or IAi
1. ii x IAIA
Gametes i i x IA IA
Fertilization IAi, IAi, IAi, IAi,
Phenotypes - All of blood group A

2. ii x IAi
Gametes i i x IA i
Fertilization iIA ii iIA IAi, ii, IAi, ii
Phenotypes Blood groups A, O, A, O
From these crossings, a man of blood group O cannot produce a child in blood group AB, but
children in blood groups A and O. Hence, it is highly unlikely that the child of blood group AB
is his child.

2) Two men in blood groups A and O respectively were in dispute over a child in blood group
AB produced from a mother of blood group B. Who is likely to be the father of the child out of
the two men?
Solution:
Blood groups genotypes
1st Man - A IAIA or IAi
2nd Man-O ii
Child - AB IAIB
Mother – B IBIB or IBi

Possible crossings of 1st man:

Phenotypes AxB
Genotypes IAIA or IAi x IBIB or IBi

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i) Gametes IA IA x IB IB
Fertilization IAIB IAIB IAIB IAIB
Phenotypes AB AB AB AB
ii) Genotype IAi x IBi
Gametes IA i x IB i
Fertilization IAIB IAi IBi ii
Phenotypes AB, A, B, O

2nd Man OxB

Genotype ii x IBIB or IBi
i) Genotype ii x IBIB
Gametes i i x IB IB
Fertilization IBi, IBi, IBi, IBi,
Phenotype All blood group B
ii) Genotype ii x IBi
Gamete i i x IB i
Fertilization IBi, ii, IBi, ii
Phenotype B O B O
From the above cross, it was found that the first is the father of the child.

Example 3.
Both Ibrahim and Halima have normal color vision. After 10 years of marriage to Ibrahim,
Halima gave birth to a color-blind daughter. Ibrahim filed for divorce, claiming he is not the
father of the child. Is Ibrahim justified in his claim of nonpaternity? Explain why. If Halima had
given birth to a color-blind son, would Ibrahim be justified in claiming nonpaternity?
Solution
Halima can either be XX or XcX
Ibrahim is XY

Genotype XX x XY
Gametes X X x X Y

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Fertilization XX, XY, XX, XY
Phenotype all homozygous in normal vision
Or
Genotype XcX x XY
Gametes Xc X x X Y
Fertilization XcX, XcY, XX, XY
Phenotype 3:1
Answer, Based on the above cross, Ibrahim cannot give birth to color blind daughter but can give
birth to color blind son.
Example 4.
Color blindness in humans is most commonly due to an X-linked recessive allele. Khadija has
normal vision, but her mother is color blind. Salim is color blind. If Khadija and Salim marry and
have a child together, what is the probability that the child will be color blind?
Solution
Color blind Mother is XcXc
Khadija is heterozygous normal XcX
Salim XcY

Genotype XcX x XcY

Gamates Xc X x Xc Y
Fertilization XcXc, XcY, XXc XY
Phenotype 2:2
The probability of having a color blind child is 50:50

GENETIC VARIATION
These are changes in characters or traits that can be found amongst various species of organisms.
These variations can either be heritable or non-heritable.
The heritable characters or variations are those that can be passed through the germ line
(sperms or eggs) during the process of sexual reproduction or through attributes passed down
through the stomata openings such as leaves, stems and roots that occasionally serve as
reproductive units (asexual reproduction). In sexually-reproducing organisms, the sperm, pollen

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and eggs constitute the germ line containing the attributes in parents which are subsequently
passed on to the offspring. Thus, the germ lines are the bearers of information that controls the
development of the offspring in the direction of the information content of the zygote which
results from the union of the sperm and the egg.
Developmental errors that do not involve the germ line may be manifested in the parents, but are
not, in general transmissible to the offspring. Such errors and post-natal accidents are referred to
as non-heritable characters e.g. amputation of limbs, or blindness that results from accidents,
weight lifting, a blacksmith or professional skills. A weight lifter or a blacksmith develop extra
muscles but this muscle development cannot be passed on to the offspring unless the offspring is
subjected to the same physical exercises as the parents.
Heritable variations may either be continuous or discontinuous
1 Continuous or indiscrete variations
These are variations that have intermediary values, which are not exact, i.e. indiscrete.
Such variations are mainly morphological or physical variations. Examples are height, weight,
length, skin pigmentation (colour), colour of eyes, fingerprints, etc. In continuous variation there
is a complete range of measurements from one extreme to the other. Height is an example of
continuous variation individuals can have a complete range of heights like 1.6, 1.61, 1.62, 1.65m.
Discontinuous or discrete variations
These are variations that do not have intermediary values. That is, the values are exact or
discrete. Such variations are mainly physiological variations. Examples are sickling character or
trait, blood group, tasters, tongue rolling, haemophilia, etc. This is where individuals fall into a
number of distinct classes or categories, and is based on features that cannot be measured across
a complete range. You either have the characteristic or you don't. Blood groups are a good
example: you are either one blood group or another, you can't be in between.
Mutation
This is a sudden, random alteration in the genotype of an organism, involving qualitative or
quantitative alterations in the genetic material itself. It may involve a single gene (point
mutation) or a whole chromosome (chromosomal aberrations). Mutation may either affect the
gene number or chromosome structure.
In biology, mutations are changes to the nucleotide sequence of the genetic material of an
organism. Mutations can be caused by copying errors in the genetic material during cell division,

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by exposure to ultraviolet or ionizing radiation, chemical mutagens, or viruses, or can be induced
by the organism, itself, by cellular processes such as hyper mutation. In multicellular organisms
with dedicated reproductive cells, mutations can be subdivided into germ line mutations, which
can be passed on to descendants through the reproductive cells, and somatic mutations, which
involve cells outside the dedicated reproductive group and which are not transmitted to
descendants, usually.
Mutation is generally accepted by the scientific community as the mechanism upon which
natural selection acts, providing the advantageous new traits that survive and multiply in
offspring or disadvantageous traits that die out with weaker organisms. These changes to the
genetic make-up of an individual which cannot be accounted for by the normal processes may
(rarely) involve chromosomal mutations (e.g. Down’s Syndrome, where an individual has
trisomy (= 3 copies) of chromosome 21; they therefore have 47 chromosomes).
Classes of mutations
1. Spontaneous mutations
Many common human diseases, often devastating in their effects, are due to mutations in single
genes. Genetic diseases arise by spontaneous mutations in germ cells (egg and sperm), which are
transmitted to future generations. For example, sickle-cell anemia, which affects 1 in 500
individuals of African descent, is caused by a single missense mutation at codon 6 of the β-
globin gene; as a result of this mutation, the glutamic acid at position 6 in the normal protein is
changed to a valine in the mutant protein. This alteration has a profound effect on hemoglobin,
the oxygen-carrier protein of erythrocytes, which consists of two α-globin and two β-globin
subunits. The deoxygenated form of the mutant protein is insoluble in erythrocytes and forms
crystalline arrays. The erythrocytes of affected individuals become rigid and their transit through
capillaries is blocked, causing severe pain and tissue damage. Because the erythrocytes of
heterozygous individuals are resistant to the parasite causing malaria, which is endemic in
Africa, the mutant allele has been maintained. It is not that individuals of African descent are
more likely than others to acquire a mutation causing the sickle-cell defect, but rather the
mutation has been maintained in this population by interbreeding. Spontaneous mutation in
somatic cells (i.e., non-germline body cells) also is an important mechanism in certain human
diseases, including retinoblastoma, which is associated with retinal tumors in children.

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 2. Induced mutations
Mutagen or mutagenic agent This is any agent that brings about mutation. It may be
physical or chemical.
a) Physical mutagenic agents
These include (1) temperature and (2) radiations. The radiation may be ionized e.g. x rays and
cosmic rays or non-ionizing, e.g. ultraviolet (UV) rays.
b) Chemical mutagenic agents
(1) Reactants – These reacts with the purine and pyrimidine bases, e.g. nitrous acid,
formaldehyde.
(2) Base analogues- These mimics a pyrimidine or purine base, e.g. 5-bromo uracil (a pyrimidine
analog), 2-amino-purine (a purine analogue).
(3) acridine dyes e.g. acridine orange, acridine yellow and proflavin.
(4) Alkylating agents e.g. mustard gases
(5) Carcinogens e.g. methyl cholanthrene
(6) Acids e.g. phenols

GENETIC SYNDROMES
1. Down syndrome
This was formerly called Mongolism. It is a syndrome as it consists of a set of abnormalities. It
occurs as a result of nondisjunction (failure of human chromosomes to separate during meiotic
anaphase. People with Down syndrome have subnormal intelligence and a characteristic facial
appearance, including a folding of the eyelid reminiscent of the epicanthic fold of Asiatic people.
A karyotype of the chromosomes a person with Down syndrome shows that they have three
copies of chromosome 21, so the condition is also known as trisomy-21. Down syndrome
children have slight to severe mental retardation.
2. Turner syndrome
Persons who have Turner syndrome are female; they do not undergo puberty and their female
secondary sex characteristics remain immature: menstruation is usually absent, breast
development is slight, and pubic hair is sparse. This syndrome is seen in 1 of 3000 female births.

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Affected women are frequently short and have a low hairline, a relatively broad chest with
widely spaced nipples. Their intelligence is usually normal. Most women who have Turner
syndrome are sterile. In 1959, C. E. Ford used new techniques to study human chromosomes and
discovered that cells from a 14-year-old girl with Turner syndrome had only a single X
chromosome; this chromosome complement is usually referred to as XO with a karyotype of 45
XO. There are no known cases in which a person is missing both X chromosomes, an indication
that at least one X chromosome is necessary for human development. Presumably, embryos
missing both Xs are spontaneously aborted in the early stages of development.
Klinefelter syndrome
Persons who have Klinefelter syndrome, which occurs with a frequency of about 1 in 1000 male
births, have cells with one or more Y chromosomes and multiple X chromosomes. The cells of
most males having this condition are XXY, but cells of a few Klinefelter males are XXXY,
XXXXY, or XXYY. Persons with this condition, though male, some breast enlargement, and
reduced facial and pubic hair. They are often taller than normal and sterile; most have normal
intelligence.
Albinism
Albinism is the lack of normal pigmentation occurs in all races. A rare condition, albinism
occurs when a person inherits a recessive allele, or group of genes, for pigmentation from each
parent. In this case, production of the enzyme tyrosinase is defective. Tyrosinase is necessary to
the formation of melanin, the normal human skin pigment. Without melanin, the skin lacks
protection from the sun and is subject to premature aging and skin cancer. The eyes, too,
colorless except for the red blood vessels of the retina that show through, cannot tolerate light.
Albinos tend to squint even in normal indoor lighting and frequently have vision problems.

BIOTECHNOLOGY
Biotechnology is the manipulation of biological organisms to make products that benefit human
beings. Biotechnology contributes to such diverse areas as food production, waste disposal,
mining, and medicine. Although biotechnology has existed since ancient times, some of its most
dramatic advances have come in more recent years. Modern achievements include the transfer of

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a specific gene from one organism to another (by means of a set of genetic engineering
techniques known as transgenic); the maintenance and growth of genetically uniform plant- and
animal-cell cultures, called clones; and the fusing of different types of cells to produce beneficial
medical products such as monoclonal antibodies, which are designed to attack a specific type of
foreign substance.
The first achievements in biotechnology were in food production, occurring about 5000 BC.
Diverse strains of plants or animals were hybridized (crossed) to produce greater genetic variety.
The offspring from these crosses were then selectively bred to produce the greatest number of
desirable traits. Repeated cycles of selective breeding produced many present-day food staples.
The modern era of biotechnology had its origin in 1953 when American biochemist James
Watson and British biophysicist Francis Crick presented their double-helix model of DNA. This
was followed by Swiss microbiologist Werner Arber's discovery in the 1960s of special
enzymes, called restriction enzymes, in bacteria. These enzymes cut the DNA strands of any
organism at precise points. In 1973 American geneticist Stanley Cohen and American biochemist
Herbert Boyer removed a specific gene from one bacterium and inserted it into another using
restriction enzymes. This event marked the beginning of recombinant DNA technology,
commonly called genetic engineering. In 1977 genes from other organisms were transferred to
bacteria. This achievement eventually led to the first transfer of a human gene, which coded for a
hormone, to Escherichia coli bacteria. Although the transgenic bacteria (bacteria to which a gene
from a different species has been transferred) could not use the human hormone, they produced it
along with their own normal chemical compounds. In the 1960s an important project used
hybridization followed by selective breeding to increase food production and quality of wheat
and rice crops.
GENE MANIPULATION (GENETIC ENGINEERING)
Genetic engineering can be described as an in vitro manipulation of genes. It refers to artificial
synthesis, modification, removal, addition and repair of the genetic material (DNA) to alter the
genotype at will. Genetic engineering, recombinant DNA technology, genetic
modification/manipulation (GM) and gene splicing are terms that apply to the direct
manipulation of an organism's genes. Genetic engineering is different from traditional breeding,
where the organism's genes are manipulated indirectly; genetic engineering uses the techniques
of molecular cloning and transformation to alter the structure and characteristics of genes

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directly. Genetic engineering techniques have found some successes in numerous applications.
Some examples are in improving crop technology, the manufacture of synthetic human insulin
through the use of modified bacteria.
Five steps involved in genetic manipulation
1. Isolation of the genes of interest
2. Insertion of the genes into a transfer vector
3. Transfer of the vector to the organism to be modified
4. Transformation of the cells of the organism
5. Separation of the genetically modified organism (GMO) from those that have not been
successfully modified
Techniques of Genetic Engineering
Much before the advent of modern biotechnology, there were several practices that were
commonly followed. Breeding (hybridisation) of plants and animals to change the genotype and
phenotype of the hybrids is probably the oldest and the most widely used technique of genetic
engineering.
NEW TECHNIQUE
In the recent years, many modern techniques have been developed which have enabled scientists
to manipulate the genetic material. These include recombinant DNA technology, transformation
and transduction.
APPLICATION GENETIC ENGINEERING
Genetic engineering enables scientists to produce clones of cells or organisms that contain the
same genes.
1. Scientists use restriction enzymes to isolate a segment of deoxyribonucleic acid (DNA) that
contains a gene of interest—for example, the gene regulating insulin production.
2. A plasmid removed from a bacterium and treated with the same restriction enzyme binds with
the DNA fragment to form hybrid plasma.
3. The hybrid plasmid is re-inserted back into the bacterium, where it replicates as part of the
cell’s DNA.
4. A large number of identical daughter cells (clones) can be cultured and their gene products
extracted for human use.

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Genetic engineering, alteration of an organism's genetic, or hereditary, material to eliminate
undesirable characteristics or to produce desirable new ones. Genetic engineering is used to
increase plant and animal food production; to help dispose of industrial wastes; and to diagnose
disease, improve medical treatment, and produce vaccines and other useful drugs. Included in
genetic engineering techniques are the selective breeding of plants and animals, hybridization
(reproduction between different strains or species), and recombinant deoxyribonucleic acid
(DNA). Human insulin was first produced in the lab using recombinant (genetically engineered)
bacteria in 1978, and five years later recombinant human insulin, used to treat diabetes mellitus,
became the first biopharmaceutical on the market.
2 SELECTIVE BREEDING
The first-known genetic engineering technique, still used today, was the selective breeding of
plants and animals, usually for increased food production. In selective breeding, only those
plants or animals with desirable characteristics are chosen for further breeding. Corn has been
selectively bred for increased kernel size and number and for nutritional content for about 7,000
years. More recently, selective breeding of wheat and rice to produce higher yields has helped
supply the world's ever-increasing need for food.
3 HYBRIDIZATION
Hybridization (crossbreeding) may involve combining different strains of a species (that is,
members of the same species with different characteristics) or members of different species in an
effort to combine the most desirable characteristics of both. For at least 3,000 years, female
horses have been bred with male donkeys to produce mules, and male horses have been bred
with female donkeys to produce hinnies, for use as work animals.
Modern Gene manipulation techniques
Recombinant DNA technology is a set of techniques for recombinant genes from different
sources and transferring the recombinant DNA into other cells, where it may be expressed. Such
gene manipulation has sparked an explosion of discoveries in molecular biology and an
industrial revolution in biotechnology
Applications of Recombinant DNA technology
1. Recombinant DNA technology has been a boom for biological research, allowing
investigators to answer questions about molecular evolution, probe details of gene organization
and control, produce and catalog proteins of interest, and map eukaryotic genes.

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2. Polymerase chain reaction (PCR) is a technique for quickly amplifying DNA in vitro.
3. Exciting medical applications of recombinant DNA technology include the large-scale
production of previously scarce pharmaceutical products; the design of safer, more effective
vaccines; the development of diagnostic tests for detecting mutations that cause genetic disease;
and the ultimate prospect of curing and preventing genetic disorders caused by single defective
genes.
4. Potential agricultural benefits of genetic engineering include the design of more
productive and disease-resistant plants and animals and improvement in food quality.
Applications of Biotechnology in Development
Today biotechnology is applied in various fields. In waste management, for example,
biotechnology is used to create new biodegradable materials.
1. One such material is made from the lactic acid produced during the bacterial fermentation
of discarded corn stalks. When individual lactic acid molecules are joined chemically,
they form a material that has the properties of plastics but is biodegradable. Widespread
production of plastic from this material is expected to become more economically viable
in the future.
2. Biotechnology also has applications in the mining industry. In its natural state, copper is
found combined with other elements in the mineral chalcopyrite. The bacterium
Thiobacillus ferrooxidans can use the molecules of copper found in chalcopyrite to form
the compound copper sulfate (CuSO4), which, in turn, can be treated chemically to obtain
pure copper. This microbiological mining process is used only with low-grade ores.
Procedures have also been developed for the use of bacteria in the mining of zinc, lead,
and other metals.
3. The field of medicine employs some of the most dramatic applications in biotechnology.
One advance came in 1986 with the first significant laboratory production of factor VIII,
a blood clotting protein that is not produced, or has greatly reduced activity, in people
who have hemophilia. As a result of this condition, hemophiliacs are at risk of bleeding
to death after suffering minor cuts or bruises. In this biotechnological procedure, the
human gene that codes for the blood-clotting protein is transferred to hamster cells grown
in tissue culture, which then produce factor VIII for use by hemophiliacs. Factor VIII was
approved for commercial production in 1992.

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