Non-Technical Paragraph: Plant reproductive structures come in many forms from fertile leaves to cones to flowers. The one commonality amongst all of these reproductive structures is the sporangium where the genetic material is halved. The transition to sporangium initiation and its subsequent development are essential for the production of plant genetic diversity, the completion of the plant life cycle, and the production of seeds. The evolution and development of sporangia are key to the success of land plants. Although the molecular genetics of sporangium development have been well studied in flowering plants, there is little comparative data across land plants. This project will investigate the molecular genetics of sporangium development in the model fern Ceratopteris. This research will allow us to understand the evolution of sporangia across plants and will fill a gap in our knowledge about plant reproduction. Furthermore, this knowledge may provide new insight in crop improvement as seed and fruit production is a major aspect of agriculture. This interdisciplinary collaborative project will train participants from high school students to postdoctoral fellows.<br/><br/>Technical Paragraph: The sporangium is the fundamental reproductive structure common to all land plants. The evolution and development of sporangia have been key to the reproductive success and diversity of land plants. The evolution of distinct sporangia evolved three times independently and was a necessary innovation for the evolution of the seed. The evolution of smaller sporangia with an effective dehiscence mechanism is key to the reproductive success of leptosporangiate ferns, the largest group of ferns. Therefore, elucidating a core sporangia development network will provide insights into not only the reproductive structure common to all plants but also the evolution and development of structures that are key to agriculture. Despite its fundamental importance, a comprehensive understanding of the evolutionary developmental genetics of the reproductive transition and concomitant sporangia development across land plants is lacking. This project will leverage the unique advantages of the model fern Ceratopteris with a combination of established and emerging technologies to address this major knowledge gap. This project will use a multi-pronged approach combining analyses of candidate gene evolution with phylogenetic, expression, and functional analyses as well as discovering new genes and the genetic network in the model fern Ceratopteris using in situ transcriptomics and LCM RNAseq. In addition, living collections will be leveraged to study the morphological diversity of reproductive structures across ferns. The collaborative project brings together unique skill sets from botany and developmental genetics to plant biotechnology. This project will further develop Ceratopteris as a model species for plant biology studies.<br/><br/>This award reflects NSF's statutory mission and has been deemed worthy of support through evaluation using the Foundation's intellectual merit and broader impacts review criteria.