Phenotypic Integration During Development and Evolution of Beetle Horns

生物体的复杂性是基因、环境条件和发育过程的产物。这些相互作用是许多生物学和医学学科的中心主题，在进化生物学中占有特别突出的地位。最终，所有生物形式和功能的进化多样化都只能通过至少其中一些相互作用的本质变化来实现。该研究项目将探索这些相互作用的程度、性质和后果，这些相互作用在一个复杂的性状类别的起源、发展和多样化中正在作为一个新的模式系统出现在综合有机生物学中：甲虫角。角甲虫提供了令人难以置信的生物体丰富性和多样性，以及实验操作的可及性。利用遗传、分子和生化方法，本研究项目将1)确定哪些基因是角形成的基础；2)探索哪些基因使角的发育或多或少地对营养条件作出反应；3)探索基因和激素在角的发育和进化过程中如何相互作用。这项研究不仅将对发育、生理学和进化领域做出新的贡献，而且还将促进发育过程的进化与自然界中指导这种进化的行为和生态机制的整合。这个研究项目也与一些教育和推广工作紧密相连。首先，6名高中教师和3名少数民族高中生将在夏季被招募到调查者的实验室，让他们沉浸在发展、进化和生态学的研究中。教师和学生将通过现有的外联项目招募，这两个调查人员都已经加入了这些项目。二是培养至少7名青年科学家（6名本科生，1名博士后），从事发育生物学、生理学、遗传学、分子生物学和进化等学科的交叉研究。最后，拟议研究的结果将有助于课程设置，并将作为进化与发展和发育可塑性与进化两门研究生课程的范例。

The complexity of organisms is the product of genes, environmental conditions, and developmental processes. These interactions are central themes in many biological and medical disciplines, and occupy a particularly prominent position in evolutionary biology. Ultimately, all evolutionary diversification of organismal form and function is only possible through changes in the nature of at least some of these interactions. The research project will explore the degree, nature and consequences of these interactions in the origin, development, and diversification of a complex class of traits that is emerging as a new model system in integrative organismal biology: beetle horns. Horned beetles offer incredible organismal richness and diversity, as well as accessibility to experimental manipulation. Using genetic, molecular, and biochemical approaches this research project will 1) determine which genes underlie the formation of horns, 2) explore which genes allow horn development to be more or less responsive to nutritional conditions, and 3) explore how genes and hormones interact during horn development and evolution. This research will not only make a novel contribution to the fields of development, physiology, and evolution, but also facilitate an integration of the evolution of developmental processes with the behavioral and ecological mechanisms that guide such evolution in nature. This research program also interfaces tightly with several educational and outreach efforts. Firstly, six high school teachers and three minority high school students will be recruited into the investigators' laboratories during the summer to immerse them in research in development, evolution, and ecology. Teachers and students will be recruited through existing outreach programs with which both investigators are already affiliated. Secondly, this project will help train at least seven young scientists (6 undergraduate students, 1 postdoc) in intensive interdisciplinary research integrating developmental biology, physiology, genetics, molecular biology and evolution. Lastly, results from the proposed research will contribute to course curricula and will be used as exemplars in two graduate courses on Evolution and Development, and Developmental Plasticity and Evolution.