Recent human studies suggest that genetic polymorphisms allow an individual to maintain optimal cognitive functioning during sleep deprivation. If such polymorphisms were not associated with additional costs, selective pressures would allow these alleles to spread through the population such that an evolutionary alternative to sleep would emerge. To determine whether there are indeed costs associated with resiliency to sleep loss, we challenged natural allelic variants of the foraging gene (for) with either sleep deprivation or starvation. Flies with high levels of Protein Kinase G (PKG) (for R ) do not display deficits in short-term memory following 12 h of sleep deprivation. However, short-term memory is significantly disrupted when for R flies are starved overnight. In contrast, flies with low levels of PKG (for s , for s2 ) show substantial deficits in short-term memory following sleep deprivation but retain their ability to learn after 12 h of starvation. We found that for R phenotypes could be largely recapitulated in for s flies by selectively increasing the level of PKG in the α/β lobes of the mushroom bodies, a structure known to regulate both sleep and memory. Together, these data indicate that whereas the expression of for may appear to provide resilience in one environmental context, it may confer an unexpected vulnerability in other situations. Understanding how these tradeoffs confer resilience or vulnerability to specific environmental challenges may provide additional clues as to why an evolutionary alternative to sleep has not emerged.A lthough sleep is a behavioral state that is conserved across a diverse range of species, the biological functions of sleep remain unknown. Sleep deprivation (SD) has been shown to negatively impact cognition, but individual responses to sleep loss can vary significantly within a population (1). Recent studies suggest that a portion of this variability may be influenced by genetic factors (2). For example, polymorphisms for PERIOD 3 (PER3), a circadian clock gene, can predict the magnitude of cognitive impairment and sleep homeostasis in response to a night of SD in humans (2). Although these genetic contributions may attenuate impairments following SD, the tradeoffs associated with resistance to sleep loss remain unknown. Presumably, the potential costs must be substantial. Thus, it is likely that the price of protection from sleep loss that can be conferred by allelic variation in one environment may induce a cost when manifested in a different environment. To date, putative costs of resiliency to sleep loss have not been identified in humans or any model organism.foraging (for), which codes for Protein Kinase G (PKG), is maintained in wild-type populations as a genetic polymorphism that results in either higher or lower levels of PKG activity (3). The allele associated with higher levels of PKG ("rover"; for R ) results in larvae with longer foraging trails between food patches, whereas the allele associated with lower levels of PKG ("sitter"; for s ) results i...

We introduce a high-resolution adult foraging assay (AFA) that relates pre- and post-ingestive walking behavior to individual instances of food consumption. We explore the utility of the AFA by taking advantage of established rover and sitter strains known to differ in a number of feeding-related traits. The AFA allows us to effectively distinguish locomotor behavior in Fed and Food-Deprived (FD) rover and sitter foragers. We found that rovers exhibit more exploratory behavior into the center of an arena containing sucrose drops compared to sitters who hug the edges of the arena and exhibit thigmotaxic behavior. Rovers also discover and ingest more sucrose drops than sitters. Sitters become more exploratory with increasing durations of food deprivation and the number of ingestion events also increases progressively with prolonged fasting for both strains. AFA results are matched by strain differences in sucrose responsiveness, starvation resistance, and lipid levels, suggesting that under the same feeding condition, rovers are more motivated to forage than sitters. These findings demonstrate the AFA's ability to effectively discriminate movement and food ingestion patterns of different strains and feeding treatments.

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Metabolism, growth, and development are intrinsically linked, and their coordination is dependent upon inter-organ communication mediated by anabolic, catabolic, and steroid hormones. In Drosophila melanogaster, the corpora cardiaca (CC) influences metabolic homeostasis through adipokinetic hormone (AKH) signaling. AKH has glucagon-like properties and is evolutionarily conserved in mammals as the gonadotropin-releasing hormone, but its role in insect development is unknown. Here we report that AKH signaling alters larval development in a nutrient stress-dependent manner. This activity is regulated by the locus dg2, which encodes a cGMP-dependent protein kinase (PKG). CC-specific downregulation of dg2 expression delayed the developmental transition from larval to pupal life, and altered adult metabolism and behavior. These developmental effects were AKH-dependent, and were observed only in flies that experienced low nutrient stress during larval development. Calcium-mediated vesicle exocytosis regulates ecdysteroid secretion from the prothoracic gland (PG), and we found that AKH signaling increased cytosolic free calcium levels in the PG. We identified a novel pathway through which PKG acts in the CC to communicate metabolic information to the PG via AKH signaling. AKH signaling provides a means whereby larval nutrient stress can alter developmental trajectories into adulthood.

Reproductive division of labor is a defining characteristic of eusociality in insect societies. The task of reproduction is performed by the fertile males and queens of the colony, while the non-fertile female worker caste performs all other tasks related to colony upkeep, foraging and nest defence. Division of labor, or polyethism, within the worker caste is organized such that specific tasks are performed by discrete groups of individuals. Ordinarily, workers of one group will not participate in the tasks of other groups making the groups of workers behaviorally distinct. In some eusocial species, this has led to the evolution of a remarkable diversity of subcaste morphologies within the worker caste, and a division of labor amongst the subcastes. This caste polyethism is best represented in many species of ants where a smaller-bodied minor subcaste typically performs foraging duties while larger individuals of the major subcaste are tasked with nest defence. Recent work suggests that polyethism in the worker caste is influenced by an evolutionarily conserved, yet diversely regulated, gene called foraging (for), which encodes a cGMP-dependent protein kinase (PKG). Additionally, flexibility in the activity of this enzyme allows for workers from one task group to assist the workers of other task groups in times of need during the colony's life.In a recent article, Lucas and Sokolowski1 report that PKG mediates behavioral flexibility in the minor and major worker subcastes of the ant Pheidole pallidula. By changing the task-specific stimulus (a mealworm to induce foraging or alien intruders to induce defensive behavior) or pharmacologically manipulating PKG activity, they are able to alter the behavior of both subcastes. They also show differences in the spatial localization of the FOR protein in minor and major brains. Furthermore, manipulation of ppfor activity levels in the brain alters the behavior of both P. pallidula subcastes. The foraging gene is thus emerging as a major player in regulating the flexibility of responses to environmental change.