13-分子生物学10-22--核小体.ppt

* * * * * * * Nucleosomes Are Covalently Modified Acetylation during replication occurs on specific sites on histones before they are incorporated into nucleosomes Nucleosomes Are Covalently Modified Acetylation associated with gene activation occurs by directly modifying specific sites on histones that are already incorporated into nucleosomes DNA Structure Varies on the Nucleosomal Surface DNA is wrapped 1.67 times around the histone octamer. DNA on the nucleosome shows regions of smooth curvature and regions of abrupt kinks. The structure of the DNA is altered so that it has an increased number of base pairs/turn in the middle, but a decreased number at the ends. Structures from Protein Data Bank: 1P34. U. M. Muthurajan, et al., EMBO J. 23 (2004): 260-271. The Path of Nucleosomes in the Chromatin Fiber 10 nm chromatin fibers consist of a string of nucleosomes. 30 nm fibers have six nucleosomes/turn, which are organized into a two-start helix. Histone H1, histone tails, and increased ionic strength all promote the formation of the 30 nm fiber. The Path of Nucleosomes in the Chromatin Fiber The 10 nm fiber is a continuous string of nucleosomes The 30 nm fiber is a two start helix consisting of two rows of nucleosomes coiled into a solenoid Levels of chromatin packaging Replication of Chromatin Requires Assembly of Nucleosomes Histone octamers are not conserved during replication, but H2A-H2B dimers and H32-H42 tetramers are conserved. There are different pathways for the assembly of nucleosomes during replication and independently of replication. Replication of Chromatin Requires Assembly of Nucleosomes Nucleosome assembly in vivo Nucleosomes Are Displaced and Reassembled During Transcription Most transcribed genes retain a nucleosomal structure, though the organization of the chromatin changes during transcription. Some heavily transcribed genes appear to be exceptional cases that are devoid of nucleosomes. RNA polymerase displaces histone octamers during tran