Development and Evolution of Beetle Horns

项目摘要：两项拟议的研究考察了甲虫中最多样化的表型特征之一：角和角状结构的发育起源和随后的进化多样性。具体地说，这项拟议的研究有三个主要目标：(1)首先，它将完成甲虫角发育模式的基因网络的首次表征。(2)其次，将采用比较的方法，探讨角发育调控的修改如何在三个不同的水平上促进角甲虫的戏剧性多样性：(I)物种内可选的有角和无角雄性形态，(Ii)性二态，以及(Iii)角的位置、大小和数量的种间多样性。(3)最后，建议的研究将开发RNA干扰方法，用于未来研究可能的角模式形成机制和角表型之间的功能关系。建议研究的内在优点和更广泛的影响：建议的研究直接解决了进化生物学中最重要的悬而未决的问题之一：新的复杂表型如何起源和多样化。同时，假设框架和实验设计牢固地建立在PI对甲虫角发育的行为生态学和进化生理学10年的研究基础上。因此，拟议的研究将对进化发育生物学领域做出新的贡献，同时促进将发育过程的进化与指导这种进化的行为、生理和生态机制相结合。最后，建议的研究旨在为未来研究甲虫角的进化和发育开发工作的RNA干扰协议。因此，拟议的研究将贡献一种以前科学家无法获得的工具，但对于未来对这些生物体的发育进化进行功能分析至关重要。拟议的研究将主要由研究生和本科生进行，并为高中生提供额外的培训和研究机会。国际和平协会坚定地致力于利用拟议的研究作为机会，继续并扩大他的倡议，将代表性不足的群体纳入他在印第安纳大学的新实验室的研究和培训。除了在同行评议的期刊上及时发表外，这项提议的结果将被用作表型可塑性进化和发展的研究生课程以及昆虫生物学的本科生课程的样本。

PROJECT SUMMARY:Two proposed research examines the developmental origins and subsequent evolutionary diversification of one of the most diverse classes of phenotypic traits: horns and horn-like structure in beetles. In particular the proposed research has three major aims: (1) First, it will complete a first characterization of the gene network that patterns horn development in beetles. (2) Secondly, using a comparative approach it will explore how modifications of the regulation of horn development have contributed to the dramatic diversification of horned beetles on three different levels: (i) alternative horned and hornless male morphs within species, (ii) sexual dimorphism, and (iii) interspecific diversity in location, size, and number of horns. (3) Lastly, the proposed research will develop RNA interference approaches for future studies of functional relationships between putative horn patterning mechanisms and horn phenotypes.INTELLECTUAL MERIT & BROADER IMPACT OF THE PROPOSED RESEARCH:The proposed research directly addresses one of the most significant unresolved questions in evolutionary biology: how do novel complex phenotypes originate and diversify. At the same time the hypotheses framework and experimental design build firmly on 10 years of research by the PI on the behavioral ecology and evolutionary physiology of horn development in beetles. The proposed research will thus make a novel contribution to the field of evolutionary developmental biology, and at the same time facilitate an integration of the evolution of developmental processes with the behavioral, physiological and ecological mechanisms that guide such evolution in nature. Lastly, the proposed research aims to develop working RNAinterference protocols for future studies on the evolution and development of beetle horns. As such the proposed research will contribute a tool previously unavailable to scientists, yet paramount for future functional analyses of developmental evolution in these organisms. The proposed research will be carried out in large part by graduate and undergraduate students and provide additional training and research opportunities to high school students. The PI is strongly committed to use the proposed research as an opportunity to continue and expand his initiative to integrate underrepresented groups into research and training in his new lab at Indiana University. Apart from timely publication in peer-reviewed journals results from this proposal will be used as exemplars in a graduate course on the Evolution and Development of Phenotypic Plasticity as well as an undergraduate course in Insect Biology.