NSFDEB-NERC; Collaborative Resource; A phytochemical "tug-of-war" and its impact on organismal diversification and niche occupancy in Caryophyllales

我们提出的工作旨在了解开花植物复杂的专门化代谢性状的逐步进化，并探索这些性状的进化如何影响开花植物随后的适应和多样化。为了解决这些问题，我们将集中在显花植物目Carlellales，这是公认的非凡的适应极端环境和异常高的多样性的代谢产物来自氨基酸酪氨酸。我们的中心假设是，酪氨酸的可用性和丰富度的变化，导致了许多谱系特异性酪氨酸衍生的代谢产物的演变，反过来又深刻地影响了物种的适应性和多样性在卡莱尔目。为了验证这一假设，我们将建立一个进化框架，为卡拉目，整合转录组，基因组和metabolicdatasets，与性状进化的模式，在宏观进化尺度。具体而言，我们将：1）进行广泛的调查，以建立酪氨酸衍生的代谢性状的发生和分布acroscarbellales; 2）检查这些酪氨酸衍生的代谢性状与生物适应和多样化模式的关联;和3）确定负责这些代谢物的生物合成的进化遗传机制。在完成这项工作后，我们希望能够全面地描述卡氏藻目中酪氨酸富集代谢的程度，确定它们与卡氏藻目生物多样性模式相关的程度，并解决复杂酪氨酸衍生代谢性状背后的遗传途径的逐步进化组装。理解复杂性状的进化是生物学家面临的一个基本挑战，因为这些特征逐步进化的方式并不总是显而易见的。此外，复杂性状组装的各个阶段（如遗传、生物化学和形态学）与随后的生物多样化模式之间的联系还没有得到很好的探索，方法学方法仍处于起步阶段。最近的进展，在植物遗传学，与集成的组学规模的数据，现在提供了及时的机会，结合组装的复杂性状与谱系特异性和nichespecificorganismal多样化。特化代谢物是在某些生态和进化背景下赋予适应性能力的化学物质。在复杂的特化代谢物下，生物合成途径的逐步性质确保了它们特别易于重建复杂性的逐步进化。而特定代谢物的遗传限制性分布是解决复杂性状对生态位特异性和品系特异性生物适应和多样化的影响的基础。因此，我们的方法，使用酪氨酸丰富的特化代谢作为一个模型系统，在Carbellales，有可能导致新的和基本的见解的原因和后果的复杂性状的进化组装在各种进化尺度。

Our proposed work seeks to understand the step-wise evolution of complex specialized metabolic traits inflowering plants, and to explore how the evolution of such traits influence subsequent adaptation anddiversification in flowering plants. To address these questions, we will focus on the flowering plant orderCaryophyllales, which is well recognized for extraordinary adaptations to extreme environments andunusually high diversity of metabolites derived from the amino-acid Tyrosine. Our central hypothesis isthat changes in availability and abundance of Tyrosine in Caryophyllales has led to the evolution ofnumerous lineage-specific tyrosine-derived metabolites that in turn has profoundly influenced theadaptation and diversification of species within Caryophyllales. To test this hypothesis, we will build anevolutionary framework for Caryophyllales, that integrates transcriptomic, genomic, and metabolicdatasets, with patterns of trait evolution, on a macroevolutionary scale. Specifically, we will: 1) performan extensive survey to establish the occurrence and distribution of tyrosine-derived metabolic traits acrossCaryophyllales; 2) examine the association of these tyrosine-derived metabolic traits with organismaladaptation and diversification patterns; and 3) determine the evolutionary genetic mechanisms responsiblefor the biosynthesis of these metabolites. On completion of the proposed work, we expect to havecomprehensively described the extent of tyrosine-enriched metabolism in Caryophyllales, to have definedthe degree to which they are associated with organismal diversification patterns across Caryophyllales,and to have resolved the stepwise evolutionary assembly of the genetic pathways underlying complextyrosine-derived metabolic traits.Understanding the evolution of complex traits is a fundamental challenge for biologists, as the stepwisefashion by which such traits have evolved is not always readily apparent. Furthermore, the connectionsbetween the various stages of complex trait assembly (e.g. genetic, biochemical, and morphological) andsubsequent organismal diversification patterns are not well explored, with methodological approachesstill in their infancy. Recent advances in phylogenetics, with the integration of -omic scale data, nowprovide timely opportunities to marry the assembly of complex traits with lineage-specific and nichespecificorganismal diversification. Specialized metabolites are chemicals that confer adaptive advantagesin certain ecological and evolutionary contexts. The stepwise nature of the biosynthetic pathwaysunderlying complex specialized metabolites ensure that they are especially tractable for reconstructingstepwise evolution of complexity. While the phylogenetically restricted distributions of specializedmetabolites are fundamental to resolving the influence of complex traits on niche-specific and lineagespecificorganismal adaptation and diversification. Our approach, using the tyrosine-enriched specializedmetabolism in Caryophyllales as a model system, therefore has the potential to lead to new andfundamental insights into the causes and consequences of the evolutionary assembly of complex traits at avariety of evolutionary scales.