BANGABANDHU SHEIKH MUJIBUR RAHMAN SCIENCE &

TECHNOLOGY UNIVERSITY GOPALGANJ - 8100

AN ASSIGNMENT ON

Special Types of Chromosome

Course Title: Cytology and Cytogenetics

Course Code: BGE357

Submitted By: Submitted To:

Name: Md. Shanti Hossen Name: Dr. Md. Sarafat Ali

Student Id:18AGR165 Assistant Professor
Year:3rd
Semester:2nd
Session :2018-19 Department of Biotechnology and
Genetic Engineering
Department of Agriculture BSMRSTU, Gopalganj
BSMRSTU ,Gopalganj

Submission Date: 21.01.2023

 Introduction

Chromosomes :
A chromosome is the thread-like, gene-carrying bodies in the nucleus of a cell.
Chromosomes are composed primarily of DNA and protein. They are visible under light
microscope in a cell during metaphase stage of mitosis. Strasburger was the pioneer man
who discovered chromosomes in 1875, and the term chromosome was first coined by
Waldeyer in 1888
.
Chemical Composition of Chromosomes:
The term chromosome refers to deeply stained filamentous body formed of chromatin. The
chromatin is composed of DNA, RNA, histones and non-histones.
The purified chromatin isolated from the interphase nuclei shows the following
composition:
(i) DNA 30—40%
(ii) RNA 1—10%
(iii) Proteins 50—65%
The chemical composition of chromatin varies considerably from species to species and in
different tissues of the same species. The chemical composition of chromosomes also
differs at different stages of the cell cycle.
At metaphase chromosomes contain on an average 15-20% DNA, 10-15% RNA and 65-
75% proteins. Out of different chemical components of chromatin, DNA is the only
component, the amount of which exhibits a constant relationship with the stage of
chromosome replication and the increase or decrease in chromosome number.

Materials of chromosomes :
The chromatin material of the eukaryotic chromosomes according to its percentage of
DNA, RNA, proteins and consequently due to its staining property, has been classified into
following kinds-
1. Euchromatin: The euchromatin is the extended form of chromatin and it forms the major
portion of chromosomes. It has special affinity for basic stains and is genetically active
because its component DNA molecule synthesizes RNA molecules only in the extended
form of chromatin.
2. Heterochromatin: The heterochromatin is a condensed inter-coiled state of chromatin.
It contains 2-3 times more DNA than euchromatin.It is genetically inert as it does not
directly synthesis of RNA (transcription) and protein and is often replicated at a different
time from the rest of the DNA.

Description
Morphology of Chromosome:
The morphology of the chromosome is suited and found following seven parts in
achromosome.
(1)Centromere,
(2) chromatid
(3) secondary constriction and satellite
(4) telomere
(5) chromomer
(6) chromonema and
(7) matrix.

Fig. Simplified Structure of Chromosome

Classification of chromosome:
 Chromosome can be classified in different ways. The various criteria which are usually
used for the classification of chromosomes include the followings:
1. Position of centromere
2. Number of centromere
3. Shape at anaphase
4. Structure and appearance
5. Role in heredity (essentiality)
6. Role in sex determination and
7. Structure and function

Special types of Chromosome:

Other than usual type of chromosomes (autosomes and sex chromosomes), the eukaryotes
possess some unusual and special types of chromosomes in some body cells or at some
particular stage of their life cycle.

Special types of Chromosomes are as follows -

1. Polytene Chromsomes or Salivary Gland Chromosomes
2. Lampbrush Chromosomes
3. Prokaryotic Nucleoids.

Type 1. Polytene Chromsomes or Salivary Gland Chromosomes:

The giant chromosomes were first observed in the cells of salivary glands, gut, trachea and
other body parts of dipteran insects by E.G. Balbiani in 1881. The name polytene was
assigned to these chromosomes by Kollar. The polytene chromosomes are much larger than
the normal somatic chromosomes. How this increase in size of chromosomes is brought
about is not known.
The giant chromosomes consist of a bundle of chromonemal fibrils which arise by a series
of about 10 consecutive duplications of the initial chromonemata that increase the DNA
content about 1,000 times the DNA content of somatic cells (Fig. 9.10). Because of the
multi-stranded condition, these chromosomes are called polytene chromosomes.
The process of reduplication of strands without separation is called endoduplication. The
homologous polytene chromosomes always remain closely paired in mitotic prophase (Fig.
9.11). This is called somatic pairing and these chromosomes are thought to be in permanent
prophase.

The polytene chromosomes bear along their entire length a series of dark bands alternated
by light bands or interbands. The dark bands are narrow or broad disc shaped structures.
They are euchromatic in nature and contain large amount of DNA, small amount of RNA
and certain basic proteins. They are feulgen positive and absorb ultraviolet (UV) light of
2600 A.
The light bands or interbands are fibrillar, feulgen negative, heterochromatic regions
containing small amount of DNA. large amount of RNA and acidic proteins and they
absorb little amount of UV light.
The number, distribution and localization of discs or bands are notably similar in
homologous polytene chromosomes of Drosophila. The centromeres of all these
chromosomes fuse to form chromocentre in Drosophila. During certain developmental
stages, the single bands or adjacent bands of polytene chromosomes produce local
reversible swellings which are called ‘chromosomes puffs’ or bulbs.
The chromonemata of polytenic chromosomes give out many series of loops laterally.
These loops or rings are known as the balbiani rings and they are rich in DNA and RNA
(Fig. 9.12).

Ultrastructure of giant polytene:

It was first investigated by Beerm (poly=many, tene=strands) chromosomes: ann and Bahr
(1954), who observed numerous fine fibrils in the Balbiani rings of Chironomus and
estimated that each chromosome contains 1000 to 2000 separate strands (corresponding to
the degree of ploidy). Later Gay (1956) observed strands 200 to 500 A in diameter in
sectioned Drosophila salivary chromosomes. The individual fibres in band and interband
regions are similar in appearance, but the fibres in the bands exhibit a considerable degree
of metaphaselike folding and are much more tightly packed. Polytene chromosomes get
their name from the fact that they are formed by many parallel chromatids, often more than
a thousand strands, which do not separate from one another following duplication. Along
each chromatid strand some regions of chromatin are tightly coiled and other regions are
less coiled, with the result that polytene chromosomes appear to consist of light and dark
bands when observed under a microscope. During larval development, specific areas on
polytene chromosomes become uncoiled, formi ng localized regions called ‘pufs’. Puffs
represent regions of active RNA synthesis (transcription). In the puff individual fibres
remain continuous across the puff and they become extended as short lateral loops (Bahr,
1954). DNA is concentrated almost en tirely in the bands. Protein and RNA is also found in
puffs. Puffing is due to the uncoiling of chromosome fibres which are usually closely
folded or coiled in the dense band regions. These fibres then project in the form of loops.

Puffs and Balbiani rings:

During their initial stages of development these bands or interbands of the chromosomes
exhibit swellings or puffs. Their appearance depends on the stage of larval development. It
is probable that the metabolic activities, required for the formation of p uffs, are related to
the secretory function of the salivary glands. The formation of this is controlled by certain
specific genes and the puffs are related with the active synthesis of RNA and proteins. This
chromosomal RNA differs from the nucleolar and c ytoplasmic RNA. The RNA of puffs is
also not similar; it differs from each other in chemical composition. Some regions show
larger puffs than others. These larger puffing regions are called Balbiani rings. These rings
are formed by the lateral stretching o f loops caused by chromonemata. These loops of
chromonemata make up Balbiani rings and give the chromosome a fuzzy outlook. The
Balbiani rings are rich in DNA and mRNA, and the formation and function of the Balbiani
rings are similar to the puffs.

Function s of giant polytene chromosomes:

(1) Main function of the polytene chromosome is to carry genes which ultimately control
physiology of an organism. These genes are formed of DNA molecules. (2) Shifting of
heterochromatin in respect to euchromatin produces giant changes called position effects.
These effects cause mutatio ns in animals as well.
(3) Heterochromatic regions contain fewer genes than euchromatic parts. Production of
nucleolar material is entirely done by heterochromatin.
(4) Chromosomes also help in protein synthesis indirectly. Nucleolus contains RNA, and
this RNA serves as a means of transmission of genetic information to the cytoplasm,
leading to the formation of specific protein.

Question1.. Why are polytene chromosomes useful in genetic studies?

Answer:
Chromonema is required to form Polytene chromosomes which are formed by
chromosomal replication without nuclear division.
The giant chromosomes are also formed by the process of somatic pairing between
homologous chromosomes.
The resultant daughter chromosomes remain aligned with each other and not separated
from each other.
This is very useful for the analysis of many facets of eukaryotic interphase chromosome
organization and the genome as a whole.

Question2.How are polytene chromosomes produced?

Answer:
Chromonema is required to form Polytene chromosomes which are formed by
chromosomal replication without nuclear division.
The giant chromosomes are also formed by the process of somatic pairing between
homologous chromosomes.
The resultant daughter chromosomes remain aligned with each other and not separated
from each other.
The salivary gland cells do not undergo mitosis and die during metamorphosis.
This chromosome is symbolized by Balbiani rings or puffs which are swollen or puffy
areas in polytene chromosomes.
This chromosome is the site of mRNA synthesis and is multi- stranded in nature.

Type 2. Lampbrush Chromosomes:

Lampbrush chromosomes are special type of giant chromosomes found in the nuclei of
oocytes of many vertebrates, such as fishes, amphibians, reptiles and birds during the
prolonged diplotene stage of first meiosis. They are also found in the nucleus of Drosophila
spermatocyte. Lampbrush chromosomes were first observed by Flemming in 1882 and
given the name by Ruckert in 1892.
These chromosomes may sometime become even larger than the polynemic or polytenic
chromosomes of salivery glands of dipterans. The largest chromosomes may sometime be
as long as 1 mm in urodele amphibians. These chromosomes consist of main axis and many
fine lateral projections or loops which give them the appearance of a test tube brush or
lampbrush (Fig. 9.13).
Actually, the main axis consists of four chromatids or two bivalent chromosomes and the
chromonemeta of these chromatids give out fine loops at the lateral sides. Only the
kinetochore bears no lateral loops. Ris (1957) studied the loops with electron microscope
and suggested that the loops were integral parts of chromonemata which are extended in
the form of major coils (Fig. 9.13).
Generally one to nine loops may arise from a single chromomeral area. The size of loop
varies from an average of 9.5µ in frog to nearly 200 µ in newt. These loops probably
consist of one DNA double helix from which fibrils project which are covered with loop
matrix consisting of RNA and proteins.
Loop formation is interpreted by Gall (1958) as a reversible physiological change which is
probably non-genetic. The number of pairs of loops increases in meiosis till it reaches a
maximum in diplotene. After diplotene stage, the number of loop pairs gradually decreases
and the loops disappear at Metaphase 1.
Physiological studies indicate that the loops of lamp-brush chromosomes and balbiani rings
of polytene chromosomes are the sites of active genes.

Functions of lampbrush:
(a) Synthesis of of Lampbrush RNA: Functions of lampbrush chromosomes, chromosomes
involve synthesis of RNA and protein by their loops. RNA is synthesized only at the thin
insertion and then carried around the loops to the thick insertion. There it may be either
destroyed or released into nucleus.
(b) Formation of yolk mater ial: There are some probabilities that lampbrush chromosomes
help in the formation of certain amount of yolk material for the egg.

Question 1. Why do lampbrush chromosomes are observed during oogenesis and

what roles do they play in egg formation?
Answer:
The Loops are clearly seen in lampbrush chromosomes during meiotic prophase in oocytes
of many species.
Lampbrush Chromosomes are concerned with vitellogenesis (Yolk formation).
This is the special kind of synapsed mid prophase or the Diplotene bivalent stage
This chromosome occurs in pairs containing homologous chromosomes containing the
point of contact known as chiasmata.
It stores components needed for rapid cell divisions during the early development of the
fertilized egg.

Question 2. In which type of stage are lampbrush chromosomes observed?

Answer:
Lampbrush chromosomes occur in the Diplotene stage of the meiosis I division cycle.

Question 3. How are lampbrush chromosomes helpful?

Answer:
Lampbrush Chromosomes are concerned with vitellogenesis (Yolk formation).
The loops present in the lampbrush chromosomes contain one to several transcriptional
units which show transcription of mRNA required for the synthesis of the substances for
growth and development of meiocytes.
Some mRNAs produced by lampbrush chromosomes may be stored as informosomes
(mRNA + protein) for producing biochemicals during the early development of the
embryo.
The lateral loops are withdrawn followed by the shortening of chromosomes, after the full
development of meiocytes.
The main function is synthesis of RNA and proteins.
Lampbrush chromosome is a model useful for studying chromosome organization, genome
function and gene expression during meiotic prophase.
This also allows the visualization of the individual transcription units.

Type 3.Prokaryotic Nucleoids:

The cell of prokaryotes or akaryobionta, such as bacteria and blue green algae, are
characterized by absence of well organized nuclei. In these cases definite nuclear envelope,
as well as nucleioli, is lacking. The electron microscopic studies have revealed that central
region of bacterial cell contains feulgen positive granules.
That area has been termed nucleoid or nucleus like structure. Feulgen positive bodies
consist of only single circular molecule of DNA which is nearly 1mm long. In cross
section, the nucleoid shows 500-900 strands.
This indicates that a single DNA strand in folded forth and back several hundred times.
Further work by a number of workers has demonstrated that the network of thread consists
of single chromosome in the form of a ring. It is known that the larger part of the DNA in
bacterial nucleoid is naked and is not combined with proteins.
The nucleoid of Escherichia coli has a point of contact with plasma membrane. Besides
single major circular chromosome, Escherichia coli in fact often possesses one or more
minor chromosomes, each called a “plasmid”, which may contain 0.5 to 2% of the total
cellular DNA. It is not yet known whether plasmids are located inside or outside the
nucleoid.
In blue-green algae, these chromidia are aggregated in the centre of the cell. This also
provides clue that the true nucleus with definite nuclear membrane is probably evolved
from the primitive type of nucleus.

B-chromosomes:
Many plant (maize, etc.) and animal (such as insects and small mammals) species, besides
having autosomes and sex chromosomes possess a special category of chromosomes called
B chromosomes without obvious genetic function. These B-- chromosomes (also called
supernumerary chromosomes, accessory chromosomes, accessory fragments, etc.) usually
havea normal structure, are somewhat smaller than the autosomes and can be
predominantly, heterochromatic (many insects, maize, etc. ) or pro dominantly euchromatic
(rye). In maize, their number per cell can vary from 0 to 30 and they adversely affect,
develo pment and fertility only when occur, in large amount. In animals, the B
chromosomes disappear from the nonreproductive (somatic) tissue and are maintained only
in the celllines that lead to the reproductive organs. B chromosomes have negative
consequenc-- es for the organism, as they have deleterious effect because of abnormal
crossing over during the meiosis of animals and abnormal nucleus divisions of the
gametoophyte plants. In animals, Bchromosomes occur more frequently in females and the
basis is non disjunction. The nondisjunction of B-- chromosomes of rye plant is found to be
caused due to the presence of a heterochromatic knob at the end of long arm of
Bchromosome. The origin of the B chromosomes is uncertain. In some animals they may
be derivatives of sex chromosomes, but this is not the rule.

Conclusion:
Polytene chromosomes were first reported by E.G.Balbiani in 1881. Polytene chromosomes
are found in dipteran flies: the best understood are those
of Drosophila, Chironomus and Rhynchosciara. They are present in another group of
arthropods of the class Collembola, a protozoan group Ciliophora,
mammalian trophoblasts and antipodal, and suspensor cells in plants. In insects, they are
commonly found in the salivary glands when the cells are not dividing.
They are produced when repeated rounds of DNA replication without cell division forms a
giant chromosome. Thus polytene chromosomes form when multiple rounds of replication
produce many sister chromatids which stay fused together.
Polytene chromosomes, at interphase, are seen to have distinct thick and thin banding
patterns. These patterns were originally used to help map chromosomes, identify
small chromosome mutations, and in taxonomic identification. They are now used to study
the function of genes in transcription. Lampbrush chromosome are a special form of
chromosome found in the growing oocytes (immature eggs) of most animals, except
mammals. They were first described by Walther Flemming and Ruckert in
1882. Lampbrush chromosomes of tailed and tailless amphibians, birds and insects are
described best of all. Chromosomes transform into the lampbrush form during the diplotene
stage of meiotic prophase I due to an active transcription of many genes. They are highly
extended meiotic half-bivalents, each consisting of 2 sister chromatids. Lampbrush
chromosomes are clearly visible even in the light microscope, where they are seen to be
organized into a series of chromomeres with large chromatin loops extended laterally.
Continuous RNA transcription is required to maintain typical chromomere-loop structure
of lampbrush chromosomes. Inhibition of transcription leads to retraction of lateral loops
into chromomeres and chromosome condensation.

References:
1.Balbiani EG (1881). "Sur la structure du noyau des cellules salivaires chez les larves
de Chironomus". Zool. Anz. 4: 637–641.
2.^ Kostoff, Dontcho (1930). "Discoid Structure of the Spireme". Journal of
Heredity. 21 (7): 323–324.
3. J (1892) Zur Entwicklungsgeschichte des Ovarialeies bei Selachiern. Anat Anz 7: 107–
158.
4.Gall JG (1966) Techniques for the study of lampbrush chromosomes. In: Prescott DM
(ed) Methods in cell physiology, vol II. Academic Press, London New York, pp 37–60.