Structure, Function & Molecular

Organization of Eukaryotic
Chromosome.
Sukamal Sarkar
Palli Siksha Bhavana
Visva Bharati
 A chromosome is an organized structure of DNA and protein found in cells. It is a single
piece of coiled DNA containing many genes, regulatory elements and other nucleotide
sequences. Chromosomes also contain DNA-bound proteins, which serve to package the DNA
and control its functions.

Chromosomes vary widely between different organisms. The DNA molecule may be circular or
linear, and can be composed of 100,000 to 10,000,000,000[1] nucleotides in a long chain.
Typically, eukaryotic cells (cells with nuclei) have large linear chromosomes and prokaryotic
cells (cells without defined nuclei) have smaller circular chromosomes, although there are many
exceptions to this rule. Also, cells may contain more than one type of chromosome; for example,
mitochondria in most eukaryotes and chloroplasts in plants have their own small chromosomes.

In eukaryotes, nuclear chromosomes are packaged by proteins into a condensed structure called
chromatin. This allows the very long DNA molecules to fit into the cell nucleus. The structure of
chromosomes and chromatin varies through the cell
cycle. Chromosomes are the essential unit for cellular
division and must be replicated, divided, and passed
successfully to their daughter cells so as to ensure the
genetic diversity and survival of their progeny.
Chromosomes may exist as either duplicated or
unduplicated. Unduplicated chromosomes are single
linear strands, whereas duplicated chromosomes
(copied during synthesis phase) contain two copies
joined by a centromere.

Compaction of the duplicated chromosomes during

Diagram1.1:- Replicated and condensed
metaphase eukaryotic chromosome. (1) Chromatid
one of the two identical parts of the chromosome
after S phase. (2) Centromere the point where
the two chromatids touch, and where the
microtubules attach. (3) Short arm. (4) Long foot.
mitosis and meiosis results in the classic four-arm
structure (pictured to the right). Chromosomal
recombination plays a vital role in genetic diversity. If
these structures are manipulated incorrectly, through
processes known as chromosomal instability and
translocation, the cell may undergo mitotic catastrophe
and die, or it may unexpectedly evade apoptosis leading
to the progression of cancer.

In practice "chromosome" is a rather loosely defined term. In prokaryotes and viruses, the term
genophore is more appropriate when no chromatin is present. However, a large body of work

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uses the term chromosome regardless of chromatin content. In prokaryotes, DNA is usually
arranged as a circle, which is tightly coiled in on itself, sometimes accompanied by one or more
smaller, circular DNA molecules called plasmids. These small circular genomes are also found
in mitochondria and chloroplasts, reflecting their bacterial origins. The simplest genophores are
found in viruses: these DNA or RNA molecules are short linear or circular genophores that often
lack structural proteins.

The word chromosome comes from the Greek (chroma, colour) and (soma, body)
due to their property of being very strongly stained by particular dyes.

Eukaryotic Chromosomes
The eukaryotic genome is made up of DNA/protein complexes called chromosomes.
Despite the compaction of the DNA (deoxyribonucleic acid) with proteins, gene sequences
embedded within chromosomes must still be available for transcription by RNA (ribonucleic
acid) polymerases and all of the DNA must be capable of being copied by DNA polymerases.

Chromosomes have two main functions: to ensure that the DNA is segregated equally to
daughter nuclei at cell division, and to ensure that the integrity of the genome is maintained
and accurately replicated in each cell cycle. The elements responsible for these functions are
centromeres, telomeres and replication origins, respectively (Figure 1.1).

Chromosome size and karyotypes

The genome size in eukaryotes varies widely, from those of the yeasts (17 million base pairs)
to those of vertebrates (3000 million base pairs or more). The genomes of some plants are
huge. In Lilium longiflorum, for example, the genome consists of 300 000 million base pairs.
Similarly, the size and number of chromosomes in any particular species the karyotype
varies widely. For example, the closely related deer species the Chinese muntjac and the Indian
muntjac have a similar genome size but the former has 23 pairs of chromosomes and the latter
only three pairs of very large chromosomes. A minimum size is required for a stable eukaryotic
chromosome. Small yeast artificial chromosomes (YACs), based on Saccharomyces cerevisiae,
are stabilized by the presence of additional DNA between the centromere and telomere, and the
minimal size appears to be 50 kb. A maximum limit also exists for chromosome size. It has
been suggested that the longest chromosome arm must not be longer than half the length of the
spindle axis at telophase. The human karyotype consists of 22 pairs of autosomes and the sex
chromosomes, XX or XY. Each human chromosome contains an average of 100 million base
pairs of DNA. Aberrations of chromosome number (aneuploidy) result from errors in
chromosome segregation. The presence of only one copy (monosomy) of any autosome is
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generally not compatible with survival in humans. On the other hand, trisomy (an extra copy of
a chromosome) of chromosomes 21 (Down syndrome), 18 and 13 are found in live births in
humans. The complex developmental abnormalities of these individuals are presumed to arise
from the increase in dosage of genes carried on these chromosomes. The three chromosomes
involved in the viable trisomies have quite low densities of genes. Trisomies of other human
chromosomes of similar or even smaller size but with high gene densities are lethal during
embryonic or fetal development. Under certain conditions and following a variety of chemical
reatments, chromosomes can be seen to be banded under the light microscope. Each
chromosome band contains approximately 510 million base pairs of DNA and the banding
patterns uniquely identify each chromosome and allow abnormalities in gross chromosome
structure to be identified. Chromosome bands have also been used to examine the evolutionary
relationships between the karyotypes of closely related species, for example between humans
and other primates.

Figure 1.2: Chromatin packaging within the chromosome. The length of a DNA molecule is shortened by 10 000-fold during the formation of
a metaphase chromosome. This is brought about by levels of chromatin packing, built one upon another. The first level of packing is brought
about by the wrapping of DNA around nucleosomes to form a structure that has been likened to beads on a string. A folding or coiling of this
level of chromatin together with additional chromosomal proteins, such as linker histones, produces the 30-nm diameter chromatin fiber. It is
still not certain how chromatin is compacted beyond this stage. The diagram illustrates one particular model, known as the radial
loop/scaffold model. In this model it is envisaged that the 30-nm fibre is arranged into loops containing _100 kb of DNA. These loops are
anchored at their bases to the chromosome scaffold. Chromatin is finally condensed both by a shortening in length of the chromosome
scaffold and by a twisting of lateral loops in toward the chromosome axis/scaffold.

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Compaction of chromatin within chromosomes
Chromosomes are maximally condensed at metaphase of the cell cycle, when a typical
mammalian chromosome will be around 10000-fold shorter than if the same length of DNA were
present as a simple, naked double helix. Packing of the DNA molecule to form the metaphase
chromosome is accomplished by an ordered series of interactions with proteins (Figure1.2). The
first, and best understood, of these is the formation of nucleosomes, which compacts the DNA
helix 7-fold. Nucleosomes consist of approximately 146 bp of DNA wrapped around a protein
core made up of the histones H2A, H2B, H3 and H4 (core histones). The N-terminal tails of the
histones, modifications of which are important in determining the type of chromatin formed,
protrude from the surface of the nucleosome. The 30-nm diameter chromatin fibre, with a
packing ratio of 1 in 50, arises through the folding and/or coiling of the nucleosomes together
with the linker histone H1. The 30-nm fibre is the basic chromatin fibre in eukaryotes. The levels
of packaging beyond the 30-nm fibre are poorly understood at the molecular and biochemical
level because the hierarchical nature of chromatin structure makes these stages hard to study.
The radial loop/scaffold model and variations on it is commonly favoured. In this model, the 30-
nm fibre is formed into loops that are attached at their bases to a nonhistone protein scaffold
(Figure 1.2).

Chromosomes in mitosis:-
The final level of chromosome packaging occurs as interphase chromosomes enter mitosis or
meiosis. The extent of this final level of packing differs among eukaryotes. In yeast there may be
only a 2-fold further condensation of the chromatin, whilst a ratio of 9:1 has been calculated for
the compaction of mammalian metaphase chromosomes over their interphase counterparts.
Phosphorylation of histones H3 and H1 accompanies the condensation of chromosomes as they
enter mitosis. Shortening of the chromosome may occur through a helical coiling of the central
chromosome axis/scaffold (Figure 2). Lateral contraction may result from condensation of the
radial loops toward the chromosome scaffold through folding, twisting or coiling (Figure 2). The
most abundant proteins of the chromosome scaffold are DNA topoisomerase II (topo II) and
ScII, which is a member of the SMC (stable maintenance of chromosomes) family. These have
important roles in both chromosome condensation and in holding the sister chromatids together
until anaphase.

Chromosomes in the interphase nucleus:-

By using fluorescence in situ hybridization (FISH) with probes for individual chromosomes, it
has been shown that each chromosome occupies a distinct globular domain, or territory, within
the interphase nucleus. In addition each chromosome territory may have a preferred location

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within the nucleus. An important consequence of this is that individual DNA sequences within a
chromosome are not free to adopt random positions within the nucleus, they are constrained to
some extent by the behavior of the whole chromosome.

Figure1.3: Natural and artificial human chromosomes: spread metaphase chromosomes from a human cell line that also harbours an artificial
chromosome. In the left panel the chromosomal DNA is stained in blue and hybridized with a probe for centromeric alpha-satellite DNA (red)
from the acrocentric chromosomes nos. 13 and 21. This probe also lights up a small chromosome (arrowed). The green signal superimposed on the
red on this small chromosome demonstrates it to be derived from a yeast artificial chromosome-based artificial chromosome. The size of this
mammalian artificial chromosome (MAC) is more easily seen in the right panel where the DNA staining is shown in black and white. Note the
brightly stained heterochromatin visible at the natural centromeres of the human chromosomes. Images courtesy of Dr. B. Grimes, MRC Human
Genetics Unit, Edinburgh, UK.

Packing ratio:-
The length of DNA divided by the length into which it is packaged For example, the
shortest human chromosome contains 4.6 x 107 bp of DNA (about 10 times the genome
size of E. coli). This is equivalent to 14,000 m of extended DNA. In its most condensed
state during mitosis, the chromosome is about 2 m long. This gives a packing ratio of
7000 (14,000/2).

To achieve the overall packing ratio, DNA is not packaged directly into final structure of
chromatin. Instead, it contains several hierarchies of organization. The first level of
packing is achieved by the winding of DNA around a protein core to produce a "bead-
like" structure called a nucleosome. This gives a packing ratio of about 6. This
structure is invariant in both the euchromatin and heterochromatin of all chromosomes.
The second level of packing is the coiling of beads in a helical structure called the 30
nm fiber that is found in both interphase chromatin and mitotic chromosomes. This
structure increases the packing ratio to about 40. The final packaging occurs when the
fiber is organized in loops, scaffolds and domains that give a final packing ratio of about
1000 in interphase chromosomes and about 10,000 in mitotic chromosomes.

Eukaryotic chromosomes consist of a DNA-protein complex that is organized in a

compact manner which permits the large amount of DNA to be stored in the nucleus of
the cell. The subunit designation of the chromosome is chromatin. The fundamental unit
of chromatin is the nucleosome.

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Chromatin:-
The unit of analysis of the chromosome; chromatin reflects the general structure of the
chromosome but is not unique to any particular chromosome

Nucleosome:
Simplest packaging structure of DNA that is found in all eukaryotic chromosomes; DNA
is wrapped around an octamer of small basic proteins called histones; 146 bp is
wrapped around the core and the remaining bases link to the next nucleosome; this
structure causes negative supercoiling

The nucleosome consists of about 200 bp wrapped around a histone octamer that
contains two copies of histone proteins H2A, H2B, H3 and H4. These are known as the
core histones. Histones are basic proteins that have an affinity for DNA and are the
most abundant proteins associated with DNA. The amino acid sequence of these four
histones is conserved suggesting a similar function for all.

The length of DNA that is associated with the nucleosome unit varies between species.
But regardless of the size, two DNA components are involved. Core DNA is the DNA
that is actually associated with the histone octamer. This value is invariant and is 146
base pairs. The core DNA forms two loops around the octamer, and this permits two
regions that are 80 bp apart to be brought into close proximity. Thus, two sequences
that are far apart can interact with the same regulatory protein to control gene
expression. The DNA that is between each histone octamer is called the linker DNA
and can vary in length from 8 to 114 base pairs. This variation is species specific, but
variation in linker DNA length has also been associated with the developmental stage of
the organism or specific
regions of the genome.

The next level of organization

of the chromatin is the 30 nm
fiber. This appears to be a
solenoid structure with about 6
nucleosomes per turn. This
gives a packing ratio of 40,
which means that every 1 m
along the axis contains 40 m
of DNA. The stability of this
structure requires the presence
of the last member of the
histone gene family, histone
H1. Because experiments that
strip H1 from chromatin maintain the nucleosome, but not the 30 nm structure, it was
concluded that H1 is important for the stabilization of the 30 nm structure.

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The final level of packaging is characterized by the 700 nm structure seen in the
metaphase chromosome. The condensed piece of chromatin has a characteristic
scaffolding structure that can be detected in metaphase chromosomes. This appears to
be the result of extensive looping of the DNA in the chromosome.

The last definitions that need to be presented are eu-chromatin and heterochromatin.
When chromosomes are stained with dyes, they appear to have alternating lightly and
darkly stained regions. The lightly-stained regions are eu-chromatin and contain single-
copy, genetically-active DNA. The darkly-stained regions are heterochromatin and
contain repetitive sequences that are genetically inactive.

Centromeres and Telomeres:-

Centromeres and telomeres are two essential features of all eukaryotic chromosomes.
Each provide a unique function that is absolutely necessary for the stability of the
chromosome. Centromeres are required for the segregation of the centromere during
me iosis and mitosis, and teleomeres provide terminal stability to the chromosome and
ensure its survival.

Centromeres are those condensed regions within the chromosome that are responsible
for the accurate segregation of the replicated chromosome during mitosis and meiosis.
When chromosomes are stained they typically show a dark-stained region that is the
centromere. During mitosis, the centromere that is shared by the sister chromatids must
divide so that the chromatids can migrate to opposite poles of the cell. On the other
hand, during the first meiotic division the centromere of sister chromatids must remain
intact, whereas during meiosis II they must act as they do during mitosis. Therefore the
centromere is an important component of chromosome structure and segregation.

Within the centromere region, most species have several locations where spindle fibers
attach, and these sites consist of DNA as well as protein. The actual location where the
attachment occurs is called the kinetochore and is composed of both DNA and protein.
The DNA sequence within these regions is called CEN DNA. Because CEN DNA can
be moved from one chromosome to another and still provide the chromosome with the
ability to segregate, these sequences must not provide any other function.

Typically CEN DNA is about 120 base pairs long and consists of several sub-domains,
CDE-I, CDE-II and CDE-III. Mutations in the first two sub-domains have no effect upon
segregation, but a point mutation in the CDE-III sub-domain com pletely eliminates the
ability of the centromere to function during chromosome segregation. Therefore CDE-III
must be actively involved in the binding of the spindle fibers to the centromere.

The protein component of the kinetochore is only now being characterized. A complex
of three proteins called Cbf-III binds to normal CDE-III regions but can not bind to a
CDE-III region with a point mutation that prevents mitotic segregation. Fur thermore,
mutants of the genes encoding the Cbf-III proteins also eliminates the ability for
chromosomes to segregate during mitosis. Additional analyses of the DNA and protein

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components of the centromere are necessary to fully understand the mechanics of
chromosome segregation.

Telomeres are the region of DNA at the end of the linear eukaryotic chromosome that
are required for the replication and stability of the chromosome. McClintock recognized
their special features when she noticed, that if two chromosomes were broken in a cell,
the end of one could attach to the other and vice versa. What she never observed was
the attachment of the broken end to the end of an unbroken chromosome. Thus the
ends of broken chromosomes are sticky, whereas the normal end is not sticky,
suggesting the ends of chromosomes have unique features. Usually, but not always, the
telomeric DNA is heterochromatic and contains direct tandemly repeated sequences.
The following table shows the repeat sequences of several species. These are often of
the form (T/A)xGy where x is between 1 and 4 and y is greater than 1.

Telomere Repeat Sequences:-

Species Repeat Sequence

Arabidopsis TTTAGGG
Human TTAGGG
Oxytricha TTTTGGGG
Slime Mold TAGGG
Tetrahymena TTGGGG
Trypanosome TAGGG
Yeast (TG)1-3TG2-3

Notice that the number of TG sequences and the number of cytosines in the yeast
sequence varies. At least for yeast, it has been shown that different strains contain
different lengths of teleomeres and that the length is under genetic control.

The primary difficulty with telomeres is the replication of the lagging strand. Because
DNA synthesis requires a RNA template (that provides the free 3'-OH group) to prime
DNA replication, and this template is eventually degraded, a short single-stranded
region would be left at the end of the chromosome. This region would be susceptible to
enzymes that degrade single-stranded DNA. The result would be that the length of the
chromosome would be shortened after each division. But this is not seen.

The action of the telomerase enzymes ensure that the ends of the lagging strands are
replicated correctly. A well-studied system involves the Tetrahymena protozoa
organism. The telomeres of this organism end in the sequence 5'-TTGGGG-3'. T he
telomerase adds a series of 5'-TTGGGG-3' repeats to the ends of the lagging strand. A
hairpin occurs when unusual base pairs between guanine residues in the repeat form.
Next the RNA primer is removed, and the 5' end of the lagging strand can be used for
DNA synthesis. Ligation occurs between the finished lagging strand and the hairpin.

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Finally, the hairpin is removed at the 5'-TTGGGG-3' repeat. Thus the end of the
chromosome is faithfully replicated. The following figure shows these steps.

The Replication of Telomeres:-

Analysis of DNA Sequences in Eukaryotic Genomes:-

The technique that is used to determine the sequence complexity of any genome
involves the denaturation and renaturation of DNA. DNA is denatured by heating
which melts the H-bonds and renders the DNA single-stranded. If the DNA is rapidly
cooled, the DNA remains single-stranded. But if the DNA is allowed to cool slowly,
sequences that are complementary will find each other and eventually base pair again.
The rate at which the DNA reanneals (another term for renature) is a function of the
species from which the DNA was isolated. Below is a curve that is obtained from a
simple genome.

The Y-axis is the percent of the DNA that remains single stranded. This is expressed as
a ratio of the concentration of single-stranded DNA (C) to the total concentration of the
starting DNA (Co). The X-axis is a log-scale of the product of the initial concentration of
DNA (in moles/liter) multiplied by length of time the reaction proceeded (in seconds).
The designation for this value is Cot and is called the "Cot" value. The curve itself is
called a "Cot" curve. As can be seen the curve is rather smooth which indicates that
reannealing occurs slowing but gradually over a period of time. One particular value that
is useful is Cot , the Cot value where half of the DNA has reannealed.

Steps Involved in DNA Denaturation and Renaturation Experiments:-

1. Shear the DNA to a size of about 400 bp.

2.Denature the DNA by heating to 100oC.

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3. Slowly cool and take samples at different time intervals.
4. Determine the % single-stranded DNA at each time point.

The shape of a "Cot" curve for a given species is a function of two factors:

1. the size or complexity of the genome; and

2. the amount of repetitive DNA within the genome

If we plot the "Cot" curves of the genome of three species such as bacteriophage
lambda, E. coli and yeast we will see that they have the same shape, but the Cot of
the yeast will be largest, E. coli next and lambda smallest. Physically, the larger the
genome size the longer it will take for any one sequence to encounter its
complementary sequence in the solution. This is because two complementary
sequences must encounter each other before they can pair. The more complex the
genome, that is the more unique sequences that are available, the longer it will take for
any two complementary sequences to encounter each other and pair. Given similar
concentrations in solution, it will then take a more complex species longer to reach
Cot.

Repeated DNA sequences, DNA sequences that are found more than once in the
genome of the species, have distinctive effects on "Cot" curves. If a specific sequence
is represented twice in the genome it will have two complementary sequences to pair
with and as such will have a Cot value half as large as a sequence represented only
once in the genome.

Eukaryotic genomes actually have a wide array of sequences that are represented at
different levels of repetition. Single copy sequences are found once or a few times
in the genome. Many of the sequences which encode functional genes fall into this
class. Middle repetitive DNA are found from 10s - 1000 times in the genome.
Examples of these would include rRNA and tRNA genes and storage proteins in plants

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such as corn. Middle repetitive DNA can vary from 100-300 bp to 5000 bp and can be
dispersed throughout the genome. The most abundant sequences are found in the
highly repetitive DNA class. These sequences are found from 100,000 to 1 million
times in the genome and can range in size from a few to several hundred bases in
length. These sequences are found in regions of the chromosome such as
heterochromatin, centromeres and telomeres and tend to be arranged as a tandem
repeats. The following is an example of a tandemly repeated sequence:

ATTATA ATTATA ATTATA // ATTATA

Genomes that contain these different classes of sequences reanneal in a different

manner than genomes with only single copy sequences. Instead of having a single
smooth "Cot" curve, three distinct curves can be seen, each representing a different
repetition class. The first sequences to reanneal are the highly repetitive sequences
because so many copies of them exist in the genome, and because they have a low
sequence complexity. The second portion of the genome to reanneal is the middle
repetitive DNA, and the final portion to reanneal is the single copy DNA. The following
diagram depicts the "Cot" curve for a "typical" eukaryotic genome

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The following table gives the sequence distribution for selected species.

Species Sequence Distribution

Bacteria 99.7% Single Copy
Mouse 60% Single Copy
25% Middle Repetitive
10% Highly Repetitive
Human 70% Single Copy
13% Middle Repetitive
8% Highly Repetitive
Cotton 61% Single Copy
27% Middle Repetitive
8% Highly Repetitive
Corn 30% Single Copy
40% Middle Repetitive
20% Highly Repetitive
Wheat 10% Single Copy
83% Middle Repetitive
4% Highly Repetitive
Arabidopsis 55% Single Copy
27% Middle Repetitive
10% Highly Repetitive

Sequence Interspersion:-
Even though the genomes of higher organisms contain single copy, middle repetitive
and highly repetitive DNA sequences, these sequences are not arranged similarly in all
species. The prominent arrangement is called short period interspersion. This
arrangement is characterized by repeated sequences 100-200 bp in length interspersed
among single copy sequences that are 1000-2000 bp in length. This arrangement is
found in animals, fungi and plants.

The second type of arrangement is long-period interspersion. This is characterized by

5000 bp stretches of repeated sequences interspersed within regions of 35,000 bp of
single copy DNA. Drosophila is an example of a species with this uncommon sequence
arrangement. In both cases, the repeated sequences are usually from the middle
repetitive class. We discussed above where highly repetitive sequences are found.

Eukaryotic Chromosome Karyotype:-

Whereas bacteria only have a single chromosome, eukaryotic species have at least one
pair of chromosomes. Most have more than one pair. Another relevant point is that
eukaryotic chromosomes are detected only occur during cell division and not during all
stages of the cell cycle. They are in their most condensed form during metaphase when
the sister chromatids are attached. This is the primary stage when cytogenetic analysis
is performed.
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Each species is characterized by a karyotype. The karyotype is a description of the
number of chromosomes in the normal diploid cell, as well as their size distribution. For
example, the human chromosome has 23 pairs of chromosome, 22 somatic pairs and
one pair of sex chromosomes. One important aspect of genetic research is correlating
changes in the karyotype with changes in the phenotype of the individual.

One important aspect of genetics is correlating changes in karyotype with changes in

phenotype. For example, humans that have an extra chromosome 21 have Down's
syndrome. Insertions, deletions and changes in chromosome number can be detected
by the skilled cytogeneticist, but correlating these with specific phenotypes is difficult.

The first discriminating parameter when developing a karyotype is the size and number
of the chromosomes. Although this is useful, it does not provide enough detail to be
begin the development of a correlation between structure and function (phenotype). To
further distinguish among chromosomes, they are treated with a dye that stains the
DNA in a reproducible manner. After staining, some of the regions are lightly stained
and others are heavily stained. As described above, the lightly stained regions are
called euchromatin, and the dark stained region is called heterochromatin. The
current dye of chose is the Giemsa stain, and the resulting pattern is called the G-
banding pattern.

C-Value Paradox:-

In addition to describing the genome of an organism by its number of chromosomes, it

is also described by the amount of DNA in a haploid cell. This is usually expressed as
the amount of DNA per haploid cell (usually expressed as picograms) or the number of
kilobases per haploid cell and is called the C value. One immediate feature of
eukaryotic organisms highlights a specific anomaly that was detected early in molecular
research. Even though eukaryotic organisms appear to have 2-10 times as many genes
as prokaryotes, they have many orders of magnitude more DNA in the cell.
Furthermore, the amount of DNA per genome is correlated not with the presumed
evolutionary complexity of a species. This is stated as the C value paradox: the
amount of DNA in the haploid cell of an organism is not related to its evolutionary
complexity. (Another important point to keep in mind is that there is no relationship
between the number of chromosomes and the presumed evolutionary complexity of an
organism.)

C Values of Organisms Used in Genetic Studies:-

Species Haploid genome

E. coli 4.5 x 103
Human 3.0 x 106
Drosophila 1.7 x 105
Maize 2.0 x 106
Aribidopsis 7.0 x 104

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A dramatic example of the range of C values can be seen in the plant kingdom where
Arabidopsis represents the low end and lily (1.0 x 10^8 kb/haploid genome) the high
end of complexity. In weight terms this is 0.07 picograms per haploid Arabidopsis
genome and 100 picograms per haploid lily genome.

Genome - the complete set of chromosomes inherited from a single parent; the
complete DNA component of an individual; the definition often excludes organelles

The classification of the chromosomes is based on the location of the

centromere.

Monocentric chromosomes (chromosomes with one centromere:-)

Usually only one centromere is present in a chromosome, and such a
chromosome is called a monocentric chromosome.

Mon centric chromosomes are of various types:-

'Metacentric chromosomes: The centromere lies in the center of the

chromosome with equal sized arms. Such chromosomes become v-shaped
during nuclear division. '

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 Sub metacentric chromosome:- The centromere is just away from the
center, resulting in unequal arm length. These chromosomes become l or
j-shaped during nuclear division.

Acrocentric chromosome: the centromere is located towards one end of

the chromosomes. One arm is very long and the other is very short.

Telocentric chromosome:- The centromere is located at one end of the

chromosome and hence has one arm. Such chromosomes are rare.

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'Dicentric or polycentric chromosomes':-

Chromosomes with two or more centromeres

Nuclear organizer chromosome: chromosomes to which the nucleolus
is usually attached.

The portion of the chromosome to which the nucleolus is attached is known as

"nuclear organizer region" or "Secondary constriction".

Sat chromosomes: sometimes a spherical chromosomal section is seen

beyond the secondary constriction. This part is known as a satellite and
such chromosomes are known as Sat chromosomes.

The filamentous part of a chromosome, i.E. The dna, as opposed to the protein,
is called the chromonema.

Basic Function of Chromosomes:-

In the broadest sense, chromosomes function is to control all the activities of a
living cell. Chromosomes are essential for the process of cell division and are
responsible for the replication, division and creation of daughter cells, that
contain correct sequences of DNA and proteins. Proteins make up one of the
most important components of the human body, they are responsible for building
muscles and tissues, growth and repair, as well as the synthesis of the
thousands of enzymes like DNA replication enzymes, produced by the body.
Protein synthesis steps, and their successful completion is the responsibility of
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genes, that are contained in chromosomes. Chromosome functions include
holding genes, the units of heredity.

Genes are located at a particular point on a chromosome, known as its locus.

Each chromosome contains DNA in a double helix structure, which houses
thousands of genes along the strand, each with their own loci. Genes are the
units that hold codes that control the building and maintenance of cells; they also
control the characteristic traits that are passed on from parents to offspring.

Chromosomes are often referred to as the 'packaging material' that hold DNA
and proteins together in eukaryotic cells (cells that have a nucleus). Cell division
is a continuous process that must occur for an organism to function, whether for
growth, repair or reproduction. During cell division stages, it is the chromosome
that is responsible for the replication and distribution of DNA amongst new cells.
Strangely enough, the functions of chromosomes in plant cells are the same as
the function of chromosomes in animal cells. In fact, chromosomes differ
structurally more between prokaryotes and eukaryotes, as opposed to different
species.

References:-

Bickmore W and Craig J (1997) Chromosome Bands: Patterns in the Genome, chap. 1, 4 and 5.
Heidelberg: Springer-Verlag.
Blow JJ (1996) Eukaryotic DNA Replication. Frontiers in Molecular Biology. Oxford: Oxford
University Press.
Greider CW (1998) Telomeres and senescence: the history, the experiment, the future. Current
Biology 8: R178R181.
Kipling D (1995) The Telomere. Oxford: Oxford University Press.
Rosenfeld MA (1997) Human artificial chromosomes get real. Nature Genetics 15: 333335.

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