Boulikas T
Institute of Molecular Medical Sciences, Palo Alto, California 94306, USA.
Int Rev Cytol. 1995;162A:279-388. doi: 10.1016/s0074-7696(08)61234-6.
Polynuceosomes are constrained into loops or domains and are insulated from the effects of chromatin structure and torsional strain from flanking domains by the cross-complexation of matrix-attached regions (MARs) and matrix proteins. MARs or SARs have an average size of 500 bp, are spaced about every 30 kb, and are control elements maintaining independent realms of gene activity. A fraction of MARs may cohabit with core origin replication (ORIs) and another fraction might cohabit with transcriptional enhancers. DNA replication, transcription, repair, splicing, and recombination seem to take place on the nuclear matrix. Classical AT-rich MARs have been proposed to anchor the core enhancers and core origins complexed with low abundancy transcription factors to the nuclear matrix via the cooperative binding to MARs of abundant classical matrix proteins (topoisomerase II, histone H1, lamins, SP120, ARBP, SATB1); this creates a unique nuclear microenvironment rich in regulatory proteins able to sustain transcription, replication, repair, and recombination. Theoretical searches and experimental data strongly support a model of activation of MARs and ORIs by transcription factors. A set of 21 characteristics are deduced or proposed for MAR/ORI sequences including their enrichment in inverted repeats, AT tracts, DNA unwinding elements, replication initiator protein sites, homooligonucleotide repeats (i.e., AAA, TTT, CCC), curved DNA, DNase I-hypersensitive sites, nucleosome-free stretches, polypurine stretches, and motifs with a potential for left-handed and triplex structures. We are establishing Banks of ORI and MAR sequences and have undertaken a large project of sequencing a large number of MARs in an effort to determine classes of DNA sequences in these regulatory elements and to understand their role at the origins of replication and transcriptional enhancers.
多聚核小体被限制形成环或结构域,并通过基质附着区域(MARs)与基质蛋白的交叉复合作用,免受染色质结构的影响以及侧翼结构域的扭转应变。MARs或SARs的平均大小为500 bp,大约每隔30 kb间隔分布,是维持基因活性独立区域的控制元件。一部分MARs可能与核心复制起点(ORIs)共存,另一部分可能与转录增强子共存。DNA复制、转录、修复、剪接和重组似乎都在核基质上发生。有人提出,经典的富含AT的MARs通过与丰富的经典基质蛋白(拓扑异构酶II、组蛋白H1、核纤层蛋白、SP120、ARBP、SATB1)协同结合到MARs上,将与低丰度转录因子复合的核心增强子和核心起点锚定到核基质上;这创造了一个富含能够维持转录、复制、修复和重组的调节蛋白的独特核微环境。理论研究和实验数据有力地支持了转录因子激活MARs和ORIs的模型。推导或提出了一组21个MAR/ORI序列的特征,包括它们在反向重复序列、富含AT序列、DNA解旋元件、复制起始蛋白位点、同聚寡核苷酸重复序列(即AAA、TTT、CCC)、弯曲DNA、DNase I超敏位点、无核小体延伸序列、多聚嘌呤延伸序列以及具有左旋和三链结构潜力的基序中的富集情况。我们正在建立ORI和MAR序列库,并开展了一个大型项目,对大量MARs进行测序,以确定这些调控元件中的DNA序列类别,并了解它们在复制起点和转录增强子中的作用。