Schematic illustration of epistasis. Two mutations A and B can interact epistatically in different ways with varying effects on fitness. The fitness of the wild-type is represented by the black baselines, and the heights of arrows represent the fitness after one mutation (WA or WB) and after both mutations (WAB). Green, positive epistasis, red, negative epistasis, black, no epistasis. In (a), two independently beneficial mutations may have their joint effect increased or diminished (WAB larger or smaller), while in (b) the independent effect of the two mutations is deleterious and beneficial, respectively, and the combined expected effect on fitness is deleterious. In (c), each mutation by itself is deleterious, but when they interact, the result can be reciprocal sign epistasis (green arrow). These sketches illustrate an additive model, where the sum of WA and WB is equal to WAB without epistasis. In our model, using the geometric mean this corresponds to taking the logarithms of the fitness.