During the past decade, tremendous progress has been made in our understanding of molecular mechanisms as well as genetic variation that underlie the adaptation of plants to fluctuating environments. It has been well established that genetic and epigenetic factors interact to create specific transcriptional responses that help plants to cope with a range of biotic and abiotic stresses. Of particular interest is the response of plants to repeated episodes of the same stress, and there is no doubt that epigenetic memory within the same generation plays an important role in acclimating or priming plants for subsequent exposure to the same stress. We are, however, only beginning to understand how such epigenetic memory is first created and then maintained. In addition, our knowledge of how and how quickly induced epigenetic changes are reset remains woefully inadequate. In AUREATE, we will use the model plant Arabidopsis to gain insights into whether and how vegetative cells, stem cells and germ cells differ in their acquisition and maintenance of epigenetic memory, using state-of-the-art methods for cell-type specific analyses. Since transposable elements are a major force shaping the distribution of epigenetic marks along the genome, and at the same time represent the most variable components of genomes, we will examine a range of genetically diverse strains in order to pinpoint the role of transposable elements in differential epigenetic memory. Next, as a first step towards understanding how plants either prevent or allow that epigenetic memory is passed on to the next generation, we will study the transmission of newly acquired epigenetic information from stem cells in the shoot apical meristem to gametes, and ultimately the soma of sexual progeny, which will be contrasted with asexual regenerants. In the course of these studies, we will have identified specific regions of the genome that are sensitive to epigenetic modification upon stress, reveal which ones are stable or reset during germline development, and which ones are stable or vary during evolution. Using genetics and targeted epigenome modification, we will then begin to reveal the underlying mechanisms responsible for epigenetic memory. In summary, the outputs of AUREATE will provide a deeper understanding of transgenerational memory in plants and its potential application to enhance stress resilience in economically important plant species.