TRANSFER OF ORGANELLE GENES TO THE NUCLEUS

细胞器基因和基因组的研究对我们的 了解真核细胞和分子生物学。的 在所有真核细胞中，细胞器的正常功能都涉及到 细胞核和细胞器遗传系统之间的精确合作。 这种功能的缺陷对人类的健康有直接的影响。 健康，从最近爆炸性的报告中可以看出， 遗传性疾病。该提案涉及两个 真核生物遗传共同进化的基本问题 细胞：l）细胞器基因如何在功能上转移到细胞 细胞核和这些基因迁移的后果是什么？ 2)细胞器的突变率能在多大程度上发生根本性的变化， 或者是整个基因组，或者是一组特定的基因， 这种变化的根本原因和后果是什么？ 首先，我们将探讨几个最近的基因转移案例， l）充分发展cox II转移作为“基因”的范例 转移当场抓获”，即，拟订一项 跨区室基因家族，它积极地筛选基因 （2）决定核与核之间的关系; 是否发生了基因转移或基因替换事件 在核糖体蛋白基因的所有三个遗传区室中， rps 7; 3）确定叶绿体基因是否转移到 核通过RNA中间体; 4）测试假设， 剪接体内含子可以从细胞器衍生的第II组进化而来 内含子;和5）确定线粒体基因组是否 从根本上说，这是必需的，也许是因为它编码了 两种关键的不可输入的呼吸蛋白 第二，我们将精确测量 天竺葵线粒体基因组的突变率， 确定其缺乏RNA编辑是否反映了大规模基因组 后处理，并调查之间可能的机械联系 这两个特点。我们还将记录 四个叶绿体特有的高度加速的进化 基因编码RNA聚合酶，以及寻找相关的 叶绿体启动子和RNA聚合酶使用的变化。

The study of organelle genes and genomes is essential to our understanding of eukaryotic cellular and molecular biology. The proper functioning of organelles in all eukaryotic cells involves a precise cooperation between nuclear and organellar genetic systems. Defects in this functioning have direct consequences for human health, as is evident from the recent explosion of reports on mitochondrially inherited diseases. This proposal addresses two fundamental questions in the genetic coevolution of the eukaryotic cell: l) How are organelle genes functionally transferred to the nucleus and what are the consequences of these genetic relocations? 2) To what extent can organelle mutation rates change radically, either across the whole genome or for a specific set of genes, and what are the underlying causes and consequences of this change? First, several cases of recent gene transfer will be explored in order to l) fully develop the coxII transfer as a paradigmatic case of "gene transfer caught in the act", i.e., the elaboration of a transcompartmental gene-family that is actively sorting out gene function between the mitochondrion and the nucleus; 2) determine whether gene transfer or gene substitution events have occurred across all three genetic compartments for the ribosomal protein gene rps7; 3) determine whether chloroplast genes are transferred to the nucleus via RNA intermediates; 4) test the hypothesis that nuclear spliceosomal introns can evolve from organelle-derived group II introns; and 5) determine whether the mitochondrial genome is fundamentally dispensable or required, perhaps because it encodes two key, unimportable respiratory proteins. Second, we will accurately measure the extraordinary elevation in mutation rate of the mitochondrial genome in Pelargonium, determine whether its lack of RNA editing reflects wholesale genome retroprocessing, and investigate a possible mechanistic link between these two features. We will also document the full extent of the highly accelerated evolution that is specific to the four chloroplast genes encoding RNA polymerase, as well as search for associated changes in chloroplast promoter and RNA polymerase usage.