Louis E J
Yeast Genetics, Institute of Molecular Medicine, John Radcliffe Hospital, Oxford, UK.
Yeast. 1995 Dec;11(16):1553-73. doi: 10.1002/yea.320111604.
Yeast chromosome ends are similar in structure and function to chromosome ends in most, if not all, eukaryotic organisms. There is a G-rich terminal repeat at the ends which is maintained by telomerase. In addition to the classical functions of protecting the end from degradation and end-to-end fusions, and completing replication, yeast telomeres have several interesting properties including: non-nucleosomal chromatin structure; transcriptional position effect variegation for genes with adjacent telomeres; nuclear peripheral localization; apparent physical clustering; non-random recombinational interactions. A number of genes have been identified that are involved in modifying one or more of these properties. These include genes involved in general DNA metabolism, chromatin structure and telomere maintenance. Adjacent to the terminal repeat is a mosaic of middle repetitive elements that exhibit a great deal of polymorphism both between individual strains and among different chromosome ends. Much of the sequence redundancy in the yeast genome is found in the sub-telomeric regions (within the last 25 kb of each end). The sub-telomeric regions are generally low in gene density, low in transcription, low in recombination, and they are late replicating. The only element which appears to be shared by all chromosome ends is part of the previously defined X element containing an ARS consensus. Most of the 'core' X elements also contain an Abf1p binding site and a URS1-like element, which may have consequences for the chromatin structure, nuclear architecture and transcription of native telomeres. Possible functions of sub-telomeric repeats include: fillers for increasing chromosome size to some minimum threshold level necessary for chromosome stability; barrier against transcriptional silencing; a suitable region for adaptive amplification of genes; secondary mechanism of telomere maintenance via recombination when telomerase activity is absent.
酵母染色体末端在结构和功能上与大多数(如果不是所有的)真核生物的染色体末端相似。末端存在富含G的末端重复序列,由端粒酶维持。除了保护末端免于降解和端对端融合以及完成复制的经典功能外,酵母端粒还有几个有趣的特性,包括:非核小体染色质结构;与相邻端粒的基因发生转录位置效应斑驳;核周边定位;明显的物理聚集;非随机重组相互作用。已经鉴定出许多参与修饰这些特性中的一种或多种的基因。这些基因包括参与一般DNA代谢、染色质结构和端粒维持的基因。在末端重复序列旁边是一个中间重复元件的镶嵌体,这些元件在不同菌株之间以及不同染色体末端之间表现出大量的多态性。酵母基因组中的许多序列冗余存在于亚端粒区域(每个末端的最后25kb内)。亚端粒区域通常基因密度低、转录水平低、重组水平低,并且复制较晚。所有染色体末端似乎唯一共有的元件是先前定义的包含ARS共有序列的X元件的一部分。大多数“核心”X元件还包含一个Abf1p结合位点和一个URS1样元件,这可能对天然端粒的染色质结构、核结构和转录产生影响。亚端粒重复序列的可能功能包括:用于将染色体大小增加到染色体稳定性所需的某个最小阈值水平的填充物;防止转录沉默的屏障;基因适应性扩增的合适区域;在端粒酶活性缺失时通过重组维持端粒的二级机制。