Nasheuer Heinz-Peter, Smith Richard, Bauerschmidt Christina, Grosse Frank, Weisshart Klaus
Institut für Molekulare Biotechnologie Abteilung Biochemie, Jena, Germany.
Prog Nucleic Acid Res Mol Biol. 2002;72:41-94. doi: 10.1016/s0079-6603(02)72067-9.
The accurate and timely duplication of the genome is a major task for eukaryotic cells. This process requires the cooperation of multiple factors to ensure the stability of the genetic information of each cell. Mutations, rearrangements, or loss of chromosomes can be detrimental to a single cell as well as to the whole organism, causing failures, disease, or death. Because of the size of eukaryotic genomes, chromosomal duplication is accomplished in a multiparallel process. In human somatic cells between 10,000 and 100,000 parallel synthesis sites are present. This raises fundamental problems for eukaryotic cells to coordinate the start of DNA replication at each origin and to prevent replication of already duplicated DNA regions. Since these general phenomena were recognized in the middle of the 20th century the regulation and mechanisms of the initiation of eukaryotic DNA replication have been intensively investigated. These studies were carried out to find the essential factors involved in the process and to determine their functions during DNA replication. These studies gave rise to a model of the organization and the coordination of DNA replication within the eukaryotic cell. The elegant experiments carried out by Rao and Johnson (1970) (1), who fused cells in different phases of the cell cycle, showed that G1 cells are competent for replication of their chromosomes, but lack a specific diffusible factor required to activate their replicaton machinery and showed that G2 cells are incompetent for DNA replication. These findings suggested that eukaryotic cells exist in two states. In G1 phase, cells are competent to initiate DNA replication, which is subsequently triggered in S phase. After completion of S phase, cells in G2 are no longer able to initiate DNA replication and they require a transition through mitosis to reenable initiation of DNA replication to take place in the next S phase. The Xenopus cell-free replication system has proved a good model system in which to study DNA replication in vitro as well as the mechanism preventing rereplication within a single cell cycle (2). Studies using this system resulted in the development of a model postulating the existence of a replication licensing factor, which binds to chromatin before the G1-S transition and which is displaced during replication (2, 3). These results were supported by genetic and biochemical experiments in Saccharomyces cerevisiae (budding yeast) and Schizosaccharomyces pombe (fission yeast) (4, 5). The investigation of cell division cycle mutants and the budding yeast origin of replication resulted in the concept of a prereplicative and a postreplicative complex of initiation proteins (6-9). These three individual concepts have recently started to merge and it has become obvious that initiation in eukaryotes is generally governed by the same ubiquitous mechanisms.
基因组的准确及时复制是真核细胞的一项主要任务。这一过程需要多种因素协同作用,以确保每个细胞遗传信息的稳定性。染色体的突变、重排或丢失对单个细胞乃至整个生物体都可能有害,会导致功能障碍、疾病或死亡。由于真核生物基因组规模庞大,染色体复制是通过多平行过程完成的。在人类体细胞中,存在10000到100000个平行合成位点。这给真核细胞带来了一些基本问题,即如何协调每个复制起点处DNA复制的起始,并防止已复制的DNA区域再次复制。自20世纪中叶人们认识到这些普遍现象以来,真核生物DNA复制起始的调控和机制一直是深入研究的对象。开展这些研究是为了找出该过程中涉及的关键因素,并确定它们在DNA复制过程中的功能。这些研究催生了一个关于真核细胞内DNA复制的组织和协调的模型。Rao和Johnson(1970年)(1)进行的巧妙实验,将处于细胞周期不同阶段的细胞进行融合,结果表明G1期细胞有能力复制其染色体,但缺乏激活其复制机制所需的一种特定的可扩散因子,并且表明G2期细胞无能力进行DNA复制。这些发现表明真核细胞存在两种状态。在G1期,细胞有能力启动DNA复制,随后在S期被触发。S期完成后,G2期细胞不再能够启动DNA复制,它们需要经历有丝分裂才能在下一个S期重新启动DNA复制。非洲爪蟾无细胞复制系统已被证明是一个很好的模型系统,可用于体外研究DNA复制以及防止在单个细胞周期内再次复制的机制(2)。使用该系统进行的研究促成了一个模型假说的提出,即存在一种复制许可因子,它在G1-S转换之前与染色质结合,并在复制过程中被取代(2,3)。酿酒酵母(芽殖酵母)和粟酒裂殖酵母(裂殖酵母)中的遗传学和生物化学实验支持了这些结果(4,