A
Focal
substitution
Past
Future
Contingency
Entrenchment
B
Positive
Epistasis
Insertions : Deletions : Point mutations
1
:
2
:
10
25700
25650
25600
25550
oe_mr
oc_mr
PE scores (unfolding
energy)
0.15
<-20
-15
-10
-5
Selection coefficient
N es
sel_d
1.0000
25750
C
N
0.9998
DOPE score
1 − (fx /fy )
1 − (fx /fy ) e
π(x → y)
P (x → y) = !
z π(x → z)
0.9996
π(x → y) =
Relative fitness
Negative
Epistasis
0.9994
wild-type
Fitness:
0.20
Sign
Epistasis
Frequency
Frequency
Reciprocal-Sign
Epistasis
0.10
DOPE score based
on homology models
0.05
Phenotype:
0.00
MKKTV…NVPGD
MKK-V…NVYGD
MKKATV…NVYGD
0.25
B
Genotype:
Fitness
A
0