I

Survival Guide for Viruses

Te dynamics of different species can be

described by the
following equation
Population of the i
thgea.es
器
⼀
fni
fitness of the Eth species

general f can depend on Ni and t

f ⼆
f.CN Nz t

When it
analyzing evolutionary dynamics
is convenient to introduce the relative
total
population xi
三
Niw_ population
From the definition its easy to see that

I
ÉE
⼆
Y Ni

The dynamics of Xi after the gauge

transformation is captured
by the
well known repeicator equation 它
 2

Replicator Equation
Lets derive the dynamics for Xdt
李器
器 表⾶ ⼆
台器 Ě器
元 f Ni 袋 Ffni
⼀
⼆

fitness of the
fxi
⼆

xiff.gr ecosystem

dxi Radiator
D TE f 4 xi equation

Note that the presence of 0 makes the replicatoreq

non linear even if all fitness f const Why
Zero sum GAME 零 和 遊戲 0 0

毙 E 等
⼆
E f Ni Efxi N

D
器 ON If 4 0the total population
is constant
 3
Ftness Landscape
⼀
important concept 它

Suppose a virus with binary genome of length

wild type 000000 f
2 ⼟⼯
mutated OIOOIO f.it
Introduce the Hamming distance between
dj
two sequences by point mutations
diidj
⼀ Mutation

Mcj
⼆
udicnufdcj matrix

One can plot the fitness fi versus the genome

Sequence
⼀
⼝ fitness landscape

relative
不
Pneation

ftnessfi

sequences
 4
Quasi species Equation
In Me presence of mutation matrix mcj the
replicator equation holds viral
no
anger The

population is described by the quasi species eq

di E 0 xi
Mijfg
⼆
在 j
mutations from sequence j
to sequence i
It is rather
interesting that the Sea can be

cast into quantum mechanical form in the

time by the
imaginary gauge transformation
hit 0⼼
出⼼ Exigent
器 冷 器 ěirxi 器 éw
⼆
ě
Efmijfxjdfxiay
器 寺 時 mij 发
⼆
 5
Introduce the Hamilton matrix Hij
⼆
⼈
Hij Mij Hcj 我

So the imaginary time Schrodinger eq.is

器 5 Hij 出
Te dynamics can be constructed from the basis

Of the eigenvectors 1 瓦 x 0 ⼯ 2 上⼯

is H
If 14 ⼼ an
eigenstates

哭 H1 ㄓ ⼆
⽀ 147
⼀

比
1 ㄓ⼼ 1比 Dé E 假

In general one can decompose the initial

state into eigenvectors

⽀ 做 度1 比 ⼝
⼆
1此

E GIED
 6
Quantum dynamics of Me eigenstates is simple

Ed É 做
1 7
to t time

1⽣ ⼼ ⼀
⽀ sē 做

In the
long time limit determine the
Population xi
1 ㄓ⼼ GET EÓ
only the ground state survives

Inverse
gauge transformation
xi ⾔出 in Because EXEI

差 ÉEI éw E 黃
We can now compute Xi from 出
合出
xi
李 ⾼出
 70
How can a virus survive

Error threshold
uckenlfl

Evolutionary Dynamics by M.A Nowak

We can use the wild

type population E in
the long time limit as

the order parameter

xi s.PE
eimt
 8

competitions between IL and EL quasi species

A EO metastable B
state
peak
mesa

tsross
ernooo
o
oto
ao o

transition CIJ.tl

When the mutation rate is nite me yet

nite the winner is net the QS
age
with highest ftness It is the QS with

reasonable tolerance will dominate in the end