Abstract
Eukaryotic cells carry a vast amount of DNA packaged in their nucleus as chromatin. During cell division, it further condenses into individual Chromosomes. The basic unit of chromatin is nucleosome which comprises 200 bp of DNA wrapped around an octamer of positively charged histone proteins (2 molecules each H2A, H2B, H3, and H4), and one molecule of Histone H1. The octamer forms a cylinder around which DNA is wrapped (∼146 bp/octamer) and continues to the similar adjacent unit (∼60 bp of linker DNA), forming a string of nucleosomes. The H1 histone brings nucleosomes closer by binding with the linker DNA. These 10–11 nm thick threads further condense into 30 nm fibres, achieving a higher level of compaction by binding with nonhistone chromosomal proteins (e.g., topoisomerase II, condensins and cohesins) to form a scaffold through which the 30 nm fibre passes, forming loops. The average size of the loops is 600 nm, each accommodating approximately 60 kb of DNA. Whereas 85% of DNA is distributed in the loops, about 15% is associated with the scaffolds. The chromatin which is condensed even at interphase is ‘heterochromatin’. Constitutive heterochromatin comprises tandem repeats of short DNA sequences and is generally devoid of genes, transcription and recombination. A potentially active part of the genome forms the ‘euchromatin’, which also has a differential distribution of DNA that shows up as mutually exclusive G- and R-bands rich in tissue-specific and housekeeping genes, highlighting a highly organised compaction of DNA in the chromatin. Compared to these bands, >100 kb long ‘topologically associating domains’ (TADs) have been identified at the genome level as the functional unit of chromatin. For its differential functional states, chromatin constantly undergoes condensation and decondensation, mediated by the post-translational modification of histones (e.g. acetylation, methylation of lysine) and the displacement of nucleosomes by an ATP-dependent complex of proteins (e.g. SWI/SNF). DNA methylation is another mode of chromatin modification that affects chromatin function. Thus, a highly dynamic system of chromatin organisation not only encapsulates a large amount of DNA in the nucleus but ensures differential gene function in an orderly fashion.
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Acknowledgement
I am grateful to my colleagues, Drs Bhagyalaxmi Mohapatra, Devanjan Sinha, and Gaurav Pandey, Banaras Hindu University, and Prof. Suresh C Jhanwar, Memorial Sloan Kettering Cancer Center, USA, and former students, Priyanka Kumari and Mukulika Ray for going through the manuscript and giving valuable suggestions. I am also grateful to that large number of students whose classroom interactions helped me understand the subject better. I also record my gratitude and appreciation to Miss Saumya Barnwal and Mr Subhankar Biswas for drawing and redrawing some of the figures for the article.
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Rajiva Raman had his university education from Banaras Hindu University. After his retirement from the Department of Zoology at BHU as Professor, he is now serving in the same department as Distinguished Professor. He is also the Senior Scientist of the Indian National Science Academy. He has a teaching experience of nearly 40 years.
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Raman, R. Chromatin is a Dynamic Structure. Reson 27, 983–1002 (2022). https://doi.org/10.1007/s12045-022-1392-4
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DOI: https://doi.org/10.1007/s12045-022-1392-4