Development, Diversity And Evolution Of Reptilian Skull Shape

在个体生活和进化过程中，是什么因素决定了动物的形状？识别塑造生物形态和生物多样性的力量被认为是21世纪最大的科学挑战之一。解决这一挑战需要大规模的跨学科方法，能够解开已知影响动物形态的众多复杂因素。脊椎动物的头骨种类繁多，由预定的发育程序和进化历史以及各种功能需求塑造而成——容纳和保护大脑和感觉器官、进食、与其他生物的互动（交流、战斗），甚至外部环境。所有这些因素共同作用来控制形状；然而，目前尚不清楚每一种对最终成年形态的贡献程度，也不清楚它们在一个谱系的进化过程中是如何发挥作用的。我雄心勃勃的提议将汇集大量的数据集、尖端的方法和世界一流的专业知识，在两个非常不同的时间尺度上精确地确定头骨形状的决定因素：在个体的发育和成长过程中，以及在进化过程中。爬行动物（包括活体和化石形式）是研究多种因素对头骨形状影响的理想模型系统：它们占据着不同的陆地、水生、海洋和空中栖息地；吃各种各样的食物；在体型上表现出巨大的差异；并拥有3亿多年前的化石记录。为了记录爬行动物头骨形状的变化，并查明推动这些变化的力量，我将采用一种全面的方法，包括：先进的3D成像和可视化的头骨和（活的爬行动物）头部软组织；当代形态计量学分析量化形式；最先进的生物力学建模技术-包括肌肉骨骼建模和有限元分析-严格确定头骨性能，如咬合力，颌闭合速度和头骨强度。这些模型将使用在项目中收集的肌肉结构和生物材料特性的新信息进行改进和验证。拟议的研究特别及时，因为它将利用大量的数据-解剖学，实验，系统发育和发育-从我自己以前的工作和正在进行的合作。整合这些不同的信息将使我能够确定不同的因素是如何相互作用形成爬行动物头骨的。这项工作将使我们比以往任何时候都更接近于理解进化生物学中最重要和最基本的问题之一：复杂的力量是如何相互作用以驱动有机体变化和生物多样性的？我将设立新的标准，在功能形态学方面进行真正的多学科综合研究，这些研究可以应用于其他有机群体和身体系统。该项目将建立新的和巩固现有的英国和海外合作，并为两名早期职业科学家提供重要的培训。不同领域的工作人员——进化和发育生物学家、古生物学家、工程师、生态学家和保护科学家——将从大量的新数据集、新的工作流程和方法进步中受益，从而激发许多未来的研究。最后，这项工作的视觉和技术方面，以及它对有魅力的活动物和化石动物的关注，将吸引公众，产生参与机会和媒体的兴趣。项目成果将被纳入教学，并在伦敦大学学院的格兰特博物馆展出，3D数字模型将免费开放，让人们更好地了解有机体是如何发展和进化的，以及不同领域的研究人员共同努力解决重大问题的重要性。

What factors determine animal shape, both during the life of an individual and during evolution? Identifying the forces shaping organismal form and biodiversity is recognised as one of the great scientific challenges of the 21st century. Addressing this challenge demands a large-scale, interdisciplinary approach capable of disentangling the numerous and complex factors known to influence animal form.Vertebrate skulls are enormously diverse, and are shaped by predetermined developmental programmes and evolutionary history, as well as by various functional demands - housing and protecting the brain and sense organs, feeding, interactions with other organisms (communication, combat), and even the external environment. All of these factors work together to control shape; however, it is unclear to what extent each contributes to final adult form or how they act during the evolution of a lineage. My ambitious proposal will bring together large datasets, cutting-edge methods, and world-class expertise to precisely identify the determinants of skull shape at two very different timescales: during the development and growth of an individual, and through evolutionary time.Reptiles (both living and fossil forms) are an ideal model system for investigating the impact of multiple factors on skull shape: they occupy diverse terrestrial, aquatic, marine and aerial habitats; consume a wide range of foods; exhibit enormous differences in body size; and feature an excellent fossil record stretching back over 300 million years. In order to document changes in reptilian skull shape and pinpoint the forces driving these changes, I will apply a holistic approach involving: advanced 3D imaging and visualization of the skull and (for living reptiles) soft tissues of the head; contemporary morphometrics analyses to quantify form; and state-of-the-art biomechanical modelling techniques - including musculoskeletal modelling and finite element analysis - to rigorously determine skull performance, such as bite forces, jaw closing speed, and skull strength. These models will be refined and validated using new information on muscle architecture and biological material properties collected during the project. The proposed research is particularly timely as it will draw on a wealth of data - anatomical, experimental, phylogenetic and developmental - from my own previous work and from ongoing collaborations. Integrating these disparate strands of information will allow me to determine how different factors interact to shape reptilian skulls.This work will bring us closer than ever to understanding one of the most important and fundamental questions in evolutionary biology: precisely how do complex forces interact with each other to drive organismal change and biodiversity? I will set new standards for performing truly multidisciplinary, comprehensive studies in functional morphology that can be applied to other organismal groups and body systems. The project will establish new and solidify existing UK and overseas collaborations, and provide vital training for two early career scientists. Workers across diverse fields - evolutionary and developmental biologists, palaeontologists, engineers, and ecologists and conservation scientists - will benefit from vast new data sets, and new workflows and methodological advances, sparking many future studies. Finally, the visual and technological aspects of this work, as well as its focus on charismatic living and fossil animals, will appeal to the general public, generating engagement opportunities and media interest. Project results will be incorporated into teaching and a museum exhibit at UCL's Grant Museum, and 3D digital models will be made freely accessible, leading to greater appreciation of how organisms develop and evolve, and the importance of researchers across different fields working together to address big questions.