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Directional selection

1

2

0.25

0.5

0.75

1

1.25

1.5

1.75

0

Number of mutations

Haploid Fitness

Beneficial

Deleterious

Multiplicative

Negative epistasis

Positive epistasis

Multiplicative

Negative epistasis

Positive epistasis

Haploid Fitness:

1

(1+s)2+e

1+s

• Spread of a beneficial mutation

• Mutation-selection balance

• Fluctuating selection (with a long period)

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