Greater effort increases perceived value in an invertebrate.

Expending effort is generally considered to be undesirable. However, both humans and vertebrates will work for a reward they could also get for free. Moreover, cues associated with high-effort rewards are preferred to low-effort associated cues. Many explanations for these counterintuitive findings have been suggested, including cognitive dissonance (self-justification) or a greater contrast in state (e.g., energy or frustration level) before and after an effort-linked reward. Here, we test whether effort expenditure also increases perceived value in ants, using both classical cue-association methods and pheromone deposition, which correlates with perceived value. In 2 separate experimental setups, we show that pheromone deposition is higher toward the reward that requires more effort: 47% more pheromone deposition was performed for rewards reached via a vertical runway (high effort) compared with ones reached via a horizontal runway (low effort), and deposition rates were 28% higher on rough (high effort) versus smooth (low effort) runways. Using traditional cue-association methods, 63% of ants trained on different surface roughness, and 70% of ants trained on different runway elevations, preferred the high-effort related cues on a Y maze. Finally, pheromone deposition to feeders requiring memorization of one path bifurcation was up to 29% higher than to an identical feeder requiring no learning. Our results suggest that effort affects value perception in ants. This effect may stem from a cognitive process, which monitors the change in a generalized hedonic state before and after reward