Research Area 4 complements RA1-3 by addressing common fundamental research questions with theoretical models. RA4 develops bioinformatics and mechanistic modeling approaches to theoretically describe and investigate intra- and intercellular processes, such as developmental transitions, resource allocation strategies, and plant-soil-microbe interactions. RA4 will advance scientific knowledge by providing alternative in silico workbenches in the form of theoretical models for the analyses of biological systems in RA1-3. The computational and mathematical frameworks will allow deeper interpretation and integration of experimental data generated within CEPLAS and beyond. Leveraging the power of coordinated experimental and theoretical plant research to derive quantitative and predictive mathematical models will provide novel insights into the emergence and evolution of complex traits and diverse interacting genetic and environmental factors and how they affect plant performance, thus enabling predictive trait manipulation. By generating new testable hypotheses, RA4 will drive the design of novel experiments to generate deeper understanding of the mechanisms interacting across different spatiotemporal scales.