The effect of coinheritance on host-symbiont coevolution

描述（由申请人提供）：拟议研究的总体目标是发展进化遗传理论，以了解宿主和微生物基因组的共同传播如何影响宿主及其共生或致病微生物之间的共同进化。我们的方法是新颖的，因为它扩展了单基因组内的核基因组合的遗传理论，跨越两个基因组，宿主和共生体的基因组合。双基因座遗传理论量化了基因组合的共遗传，无论是在相同的基因组还是在不同的基因组中：共遗传越大，两个基因的进化轨迹结合的程度就越大。我们的研究结果将使我们能够解释模式的跨基因组共变异（种间不平衡）在th潜在的因果进化过程。我们认为，共同遗传决定了宿主-共生体协会是朝着增强的毒力方向发展还是朝着良性的互利共生方向发展。当应用于核和线粒体基因组合时，我们成功地预测了从线粒体到核的功能基因转移的频率将是自交程度的函数，并发现功能基因的基因组间转移在近交物种中发生的可能性是在异交物种中的10倍。线粒体和核基因在自交交配系统中的共遗传允许选择比核基因和线粒体基因独立遗传的交配系统更有效地作用于基因组合。尽管线粒体像某些共生体一样是完全垂直传播的，但我们预计在部分垂直传播的系统或共生体在遗传亲属宿主之间传染传播的系统中也会表现出类似的效果。在这项拟议的研究中，我们将解决五个具体目标，这将发展理论，以解决1）种间不平衡的产生和维持，2）宿主共生体协会中的功能基因丢失或转移，3）传播模式，毒力和种群遗传结构的共同进化。我们建议在先前NIH资助的基础上发展这种更普遍的理论。 公共卫生相关性：本研究的总体目标是建立一个宿主和共生体基因组共同进化和共同传播的一般进化理论。这是目前人类微生物组研究及其对人类健康的解释中缺失的重要组成部分（Toft和Andersson，2011年）。在人类病原体的背景下，我们可以更好地了解疾病出现的条件以及有利于疾病毒力增加和减少的条件（安德森和May，1979; May和安德森，1979）。

DESCRIPTION (provided by applicant): The overall goal of the proposed research is to develop evolutionary genetic theory for understanding how co-transmission of host and microbial genomes affects co-evolution between hosts and their symbiotic or pathogenic microbes. Our approach is novel in that it extends theory of the inheritance of nuclear gene combinations within single genomes to gene combinations across two genomes, those of host and symbiont. Two-locus descent theory quantifies the coinheritance of gene combinations, whether in the same or in different genomes: the greater the coinheritance, the greater the degree to which the evolutionary trajectories of both genes are conjoined. Our results will allow us to interpret patterns of trans-genomic co- variation (interspecies disequilibria) in terms of th underlying causal evolutionary processes. We propose that coinheritance determines whether a host-symbiont association evolves toward enhanced virulence or toward benign mutualistic symbiosis. When applied to nuclear and mitochondrial gene combinations, we successfully predicted that the frequency of functional gene transfer from the mitochondria to the nucleus would be a function of the degree of selfing and discovered that inter-genomic transfer of functional genes is ten-fold more likely to occur in inbreeding species than in outcrossing species. Coinheritance of mitochondrial and nuclear genes in selfing mating systems allows selection to act more effectively on gene combinations than mating systems where nuclear and mitochondrial genes are inherited independently. Althoughmitochondria are wholly vertically transmitted like some symbionts, we expect similar effects to be manifest in systems with partial vertical transmission or in systems with contagious transmission of symbionts among host that are genetic relatives. In this proposed research we will address five specific aims which will develop theory to address the 1) generation and maintenance of interspecies disequilibria, 2) functional gene loss or transfer in host-symbiont associations, 3) coevolution of transmission mode, virulence, and population genetic structure. We propose to develop this more general theory, building upon progress from prior NIH funding. PUBLIC HEALTH RELEVANCE: The overall goal of our proposed research is to develop a general evolutionary theory of the co- evolution and co-transmission of host and symbiont genomes. This is an important missing component of current investigations of the human micro-biome and its interpretation in regard to human health (Toft and Andersson 2011). In the context of human pathogens, we may better understand the conditions for disease emergence as well as those that favor increases and decreases in disease virulence (Anderson and May 1979; May and Anderson, 1979).