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Dr Moni Kumari

M.Sc 2nd and MSc 4th semester

BSc 3rd year

Dated 20-05-2020

Special chromosomes

(1) Polytene chromosome

Polytene chromosomes are larges specific interphase chromosomes which have thousands
of DNA strands.

These chromosomes, first observed in cells of insect salivary glands of Drosophila.

Chromatin structure is also unusually visible in certain insect cells.


They are about 0.5 mm in length and 20 μm in diameter.
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.
Or

Polytene chromosomes are produced by endoreplication, in which


chromosomal DNA undergoes mitotic replication, but the strands do not separate.
As for example: Ten rounds of endoreplication produces 210 = 1,024 DNA strands, which
when arranged alongside of each other produce distinctive banding patterns
They are very large and display a characteristic band–interband morphology. Along the linear
axis polytene chromosome have variation in the concentration of the chromatin. Regions of
high concentrations are known as chromomeres (bands), and regions with low concentrations
are known as interchromomeres (interbands).
The pattern of bands and interbands in each polytene chromosome is specific for the species
These band patterns of Polytene Chromosome used to help map chromosomes, recognize
small chromosome mutations, and in taxonomic classification. They are now used to study
the function of genes in transcription.
The dark bands contain more DNA and less RNA. The interbands contain more RNA and
less DNA. The amount of DNA in interbands ranges from 0.8 - 25%
They provide a high level of function in certain tissues such as salivary glands.
First reported by E. G. Balbiani in 1881 in larval salivary glands, Malpighian tubules,
intestine, hypoderm and muscles of Chironomus plumosus as a cylindrical cord that
repeatedly unravelled and filled the nucleus.

Fig 1 Drak and light band of Polytene Chromosome

Fig 2 Drawing of a polytene chromosome set of Drosophila melanogaster.

Figure 2 explains: Each parental chromosome is tightly paired with its homologue (somatic
synapsis). There are regions where two homologous chromosomes are separated (asynapsis).
All the chromosomes are linked together by the pericentromeric regions to create a single
chromocentre.
The morphology of polytene chromosomes can vary widely due to the differing degrees of
synapsis of the DNA strands

Benefits of using polytene chromosomes

In addition to increasing the volume of the cells' nuclei and causing cell
expansion, polytene cells may also have a metabolic advantage as multiple copies of genes
permits a high level of gene expression.
Polyteny arises in tissues, organs and at developmental stages when there is need for the rapid
development of an organ at an unaltered high level of function.
Organs containing cells with polytene chromosomes involved in intense secretory functions.

Question: What is a puff in a polytene chromosome?

Answer: The bands of polytene chromosomes become enlarged at certain times to form
swellings called puffs. The formation of puffs is called puffing. In the regions of puffs, the
chromonemata uncoil and open out to form many loops. The puffing is caused by the
uncoiling of individual chromomeres in a band

Puffs are reversible modifications of polytene chromosomes; they originate from single bands
or from single bands and part of the adjoining interband.

These puffs and Balbiani rings are sites of active RNA synthesis; therefore, polytene
chromosomes and more specifically puffs and Balbiani rings are excellent tools to study the
induction and process of transcription

They are formed of DNA, RNA and a few proteins.

Chromonemata: 1. the coiled mass of threads visible within a nucleus at the start of cell
division. 2. a coiled chromatin thread within a single chromosome.

(B) Lampbrush Chromosome

Lampbrush chromosomes were discovered in salamander egg cells (Ambystoma


mexicanum) by Flemming in 1882.

They measure about 1500 to 2000 micrometer in length as opposed to 15–20 μm during later
stages of meiosis.
They are extremely large synapsed homologous chromosomes, seen in the diplotene stage of
prophase-I in meiosis.
Chromosomes transform into the lampbrush form during the diplotene stage of
meiotic prophase I due to an active transcription of many genes

Lampbrush chromosome 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.

LBC chromomeres can be seen in both fixed and living chromosome preparations

(Chromomere are one of the small bead-shaped and heavily staining masses of coiled
chromatin that are linearly arranged along the chromosome.)

Giant lampbrush chromosomes (LBCs) typical for growing oocytes of various animal species
are characterized by a specific chromomere-loop appearance and massive transcription.

LBC are highly decondensed because of intensive transcriptional activity.

In average, avian lampbrush chromosomes are 20–30 times longer than corresponding mitotic
metaphase chromosomes

An array of chromomeres constitutes an axis of each LBC, with neighboring chromomeres


being connected by thin decondensed chromatin threads (interchromomeric fibers).

In LBC chromomeres are unevenly distributed along the chromosome axis

Arrays of massive and prominent chromomeres alternate with regions of small and medium-
sized ones

Besides, apart from chromomeres with numerous pairs of extended loops, there are some
chromomeres lacking recognizable loops

Since chromomeres constantly appear in the same positions one can develop cytological
maps reflecting the number, size, and general pattern of distribution of chromomeres along
LBC's axes

From structural point of view LBC chromomeres are thought to represent a rosette of
microloops, which are connected by protein clips at their bases.

Longitudinal and transverse clips are generally cohesin and condensin complexes

LBC axes lack any linker histones H1

Significance of LBC:
(1) LBC are useful model for studying chromosome organization, genome function and
gene expression during meiotic prophase, since they allow the individual transcription units
to be visualized.

(2) Lampbrush chromosomes are widely used for high-resolution mapping of DNA sequences
and construction of detail cytological maps of individual chromosomes.

Question: How are Lampbrush chromosomes formed?

Answer: Chromosomes transform into the lampbrush form during the diplotene stage of
meiotic prophase I due to an active transcription of many genes

The main difference between polytene and lampbrush chromosome is that polytene
chromosomes occur in the salivary glands and other tissues of insects
whereas lampbrush chromosomes occur in the oocytes of vertebrates except for
mammals and some invertebrates.

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