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Last Update: 6 December 2017

Nucleosome Concept M - 36
In resting, non-dividing eucaryotic cells chromosomal material made up of nucleoprotein material
(nucleic acid (DNA) and proteins) called chromatin. Histones are most often as a protein. In certain cases
protamins are found instead of Histones woodcock (1973), Oudet et al. (1975). The former first showed that
under the electron microscope chromatin appears to have a string of beads structure and is made up of number
of repeating units. Oudet called them Nucleosome. The Nucleosome concept represents the latest chromosome
model of DNA Protein association. The Nucleosome are connected each other by the linker / spacer DNA.
Chromatin is amorphous and is composed of repeating structure called nucleosome. In other words
nucleosome is the structural unit of chromatin fibre and appears is "beads on a string".

STRUCTURE : Nucleosome consists of two parts -


i) Core part & ii) Connecting thread or linker DNA.
i) Core Part This nuclease-resistant structure is ellipsoidal (about
110 in diameter and 60 high) and is made up of core
proteins and core DNA.
ii) Core proteins are known as core Histones. Histones are small basic
proteins, found in all eucaryotic chromatin and are major
structural protein of chromatin, act as suppressor to gene
expression.

Histones are rich in basic amino acids like Arginine and Lysine and completely lack of tryptophan.
They are highly modified proteins. These are highly conserved proteins with very little variation in their amino
acid sequence.
There are five principal classes of Histones - H1 (H5), H2A, H2B, H3, H4 . On the basis of their relative
contents of arginine and lysine Histones are classified into three groups
i) Very lysine rich - H1 (H5)
ii) Lysine rich - H2A, H2B
iii) Arginine rich - H3 & H4.
Core Histones form histone octamer and this octamer is formed by two molecules of each of Histones
H2A, H2B, H3, H4 and is appeared to be arranged in the term of two flat tetramers.

Core DNA : It consist of 146 - nucleotide-pair length of DNA physical studies (X-ray diffraction) of
nucleosome core crystals have shown that the DNA wound as 13/4 turns of a super helix around
the outside of the histone octamer.

Linker DNA - The size of the linker DNA varies from species to species and from one cell type to
other. Linkers as short as 8 nucleotide - pairs and as long as 114 nucleotide pairs have
been reported.
After partial nuclease digestion of chromatin an approximately 200 nucleotide - pair length of DNA is
found associated with each nucleosome. So, generally the length of linker DNA is 54 bp. This linkers are
susceptible to nuclease attack.
H1 stabilizes two full turns of DNA super helix and other evidence indicates that H1 is involved in the
coiling or folding of the nucleosome fibre to form a 300 chromatin fibres.
There are some non-histone proteins, which are associated with chromatin fibre, and these are structural
enzymes and other protein. These are acidic in nature and is involved in gene expression.
Solenoid Structure :
Electron micrographs of isolated metaphase chromosomes show masses of tightly coiled or folded
lumpy fibres. These chromatin fibres have an average diameter of 300 . H1 is involved in this super coiling
of the 100 nucleosome to produce 300 chromatin fibre.

Fig. 1 Structure of nucleosome.

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