Reeves R
Biochim Biophys Acta. 1984 Sep 10;782(4):343-93. doi: 10.1016/0167-4781(84)90044-7.
Eukaryotic chromatin has a dynamic, complex hierarchical structure. Active gene transcription takes place on only a small proportion of it at a time. While many workers have tried to characterize active chromatin, we are still far from understanding all the biochemical, morphological and compositional features that distinguish it from inactive nuclear material. Active genes are apparently packaged in an altered nucleosome structure and are associated with domains of chromatin that are less condensed or more open than inactive domains. Active genes are more sensitive to nuclease digestions and probably contain specific nonhistone proteins which may establish and/or maintain the active state. Variant or modified histones as well as altered configurations or modifications of the DNA itself may likewise be involved. Practically nothing is known about the mechanisms that control these nuclear characteristics. However, controlled accessibility to regions of chromatin and specific sequences of DNA may be one of the primary regulatory mechanisms by which higher cells establish potentially active chromatin domains. Another control mechanism may be compartmentalization of active chromatin to certain regions within the nucleus, perhaps to the nuclear matrix. Topological constraints and DNA supercoiling may influence the active regions of chromatin and be involved in eukaryotic genomic functions. Further, the chromatin structure of various DNA regulatory sequences, such as promoters, terminators and enhancers, appears to partially regulate transcriptional activity.
真核染色质具有动态、复杂的层次结构。活跃基因转录一次仅在其中一小部分上发生。虽然许多研究人员试图对活跃染色质进行表征,但我们距离了解所有将其与非活性核物质区分开来的生化、形态和组成特征仍有很大差距。活跃基因显然包装在改变的核小体结构中,并与比非活性结构域凝聚程度更低或更开放的染色质结构域相关。活跃基因对核酸酶消化更敏感,可能含有特定的非组蛋白,这些非组蛋白可能建立和/或维持活跃状态。变体或修饰的组蛋白以及DNA本身改变的构型或修饰同样可能参与其中。关于控制这些核特征的机制实际上知之甚少。然而,对染色质区域和特定DNA序列的可控可及性可能是高等细胞建立潜在活跃染色质结构域的主要调控机制之一。另一种控制机制可能是将活跃染色质分隔到细胞核内的某些区域,也许是核基质。拓扑限制和DNA超螺旋可能影响染色质的活跃区域,并参与真核基因组功能。此外,各种DNA调控序列,如启动子、终止子和增强子的染色质结构似乎部分调节转录活性。