Living organisms may acclimate to environmental changes through epigenetic modifications to DNA, which alter the way genetic instructions are interpreted without altering the DNA code itself. While these modifications to organismal phenotype or function can be reversible, some of them may be inherited by offspring, potentially producing multiple, heritable outcomes from a single genome and affecting ecological and evolutionary outcomes. This project uses symbiotic, metabolically complex reef building corals as a model system to test the connections between physiological, epigenetic, and metabolic states, and predict how population and community dynamics are influenced by epigenetically-modulated phenotypes. This work will advance biological knowledge by delineating fundamental links (Rules of Life) between ubiquitous organismal energetic processes, epigenetics, and eco-evolutionary outcomes. The Broader Impacts activities parallel the project's integrative approach, linking insights from Environment x Energetics x Epigenetics x Ecology for Education into an E5 platform. The E5 platform will provide i) early career STEM training, ii) local and global community education, and iii) educational resources for open science, quantitative approaches, and research reproducibility. Further, this E5 platform will train and inform the next generation of diverse scientists and public by combining local and global initiatives focusing on groups underrepresented in STEM. This project examines how nutrient metabolism in the mitochondria generates cofactors and energy that will instruct the epigenetic machinery in the cell nucleus to modulate genome function to appropriately respond to environmental conditions. Environmentally-responsive metabolic function and energetic-epigenetic linkages act as drivers of complex emergent phenotypes. To elucidate relationships that are the basis for Rules of Life with respect to epigenetics, this project will use integrative experimental and modeling approaches focused on reef building corals to: 1) link nutritionally-provisioned metabolites with epigenetic and organismal state through seasonal sampling across environmental gradients; 2) expand current Dynamic Energy Budget (DEB) models for symbiotic organisms to further integrate critical facets of nutritional symbiosis and calcification; 3) experimentally modulate metabolic and therefore epigenetic states through repeated exposure to increased temperature and nutrients, to test intra- and trans-generational epigenetic inheritance; 4) use DEB theory to identify shifts in energetics associated with epigenetic modulation, and link these sub-organismal processes to higher levels of organization; and 5) integrate findings into a generalizable, predictive eco-evolutionary model that links nutritional interactions, metabolic states, and subsequent epigenetic effects to the timescales regulating organismal processes and eco-evolutionary outcomes. This effort will provide characterization of environmental epigenetic phenomena in ecosystem-engineering marine invertebrates. This characterization includes determining the mechanisms and the degree of epigenetic 'memory' both within and across generations. By including information on environmental legacies, propagated by epigenetics, this project will advance both organismal and population-based models and improve capacity to predict responses to acute and chronic environmental signals.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.