Early life adversity has known impacts on adult health and behavior, yet little is known about the gene-environment interactions (GEIs) that underlie these consequences. We used the fruit fly Drosophila melanogaster to show that chronic early nutritional adversity interacts with rover and sitter allelic variants of foraging (for) to affect adult exploratory behavior, a phenotype that is critical for foraging, and reproductive fitness. Chronic nutritional adversity during adulthood did not affect rover or sitter adult exploratory behavior; however, early nutritional adversity in the larval period increased sitter but not rover adult exploratory behavior. Increasing for gene expression in the mushroom bodies, an important center of integration in the fly brain, changed the amount of exploratory behavior exhibited by sitter adults when they did not experience early nutritional adversity but had no effect in sitters that experienced early nutritional adversity. Manipulation of the larval nutritional environment also affected adult reproductive output of sitters but not rovers, indicating GEIs on fitness itself. The natural for variants are an excellent model to examine how GEIs underlie the biological embedding of early experience.genotype-environment interaction | plasticity T he question of how individual differences arise is fundamental to biology, psychology, and precision medicine (1, 2). From studies on mammals, we know that early adversity places individuals on developmental trajectories for health and behavior that can last a lifetime (3, 4). However, for the most part, this idea has not been investigated in simple model genetic organisms, such as the worm Caenorhabditis elegans or the fruit fly Drosophila melanogaster, in which gene manipulation can be readily accomplished. Two biological mechanisms that can underlie individual differences, and that go beyond the obsolete notion of nature or nurture, are gene-environment interactions (GEIs) and epigenetics. Here, we explore GEIs in the context of early adversity. In particular, we address how early adversity interacts with natural variants of the foraging (for) gene to affect adult behavior and fitness in D. melanogaster.Within human populations, nutritional adversity can lead to substantive effects on cognitive and behavioral development (5, 6). The question of how early adversities perturb normal development is a difficult one, because both the strength and timing of adversity can matter. For example, in humans, detrimental effects are seen after chronic protein or carbohydrate deprivation, yet one or a few acute deprivations have little long-term effect (7). Similarly, in fruit flies, levels of hemolymph carbohydrate affected by 3 h of food deprivation return to normal levels after just 2 h of refeeding (8). Moreover, not all individuals are affected equally by early nutritional adversity (9, 10), and these individual differences arise from GEIs.In evolutionary biology, GEIs can be important for the maintenance and expression of both phenotypic plasticit...