The role of filament recycling in stable flows of 2D active networks

Committee meeting

William McFadden 2016

How to change your own shape

how can an organism rearrange itself internally and externally?

Don Kane (U. Rochester)

Cellular processes are the drivers of macroscopic reorganization

Yumura et al. 2013

 

ed munro

crawling

division

polarization

 cellular reorganization via cortical flows

Mmayer et al. 2010

 

ed munro

flows of cortical actomyosin cytoskeleton

actin filaments

and myosin motors

2D disordered network

flows of cortical actomyosin cytoskeleton

Actomyosin networks generate forces and spontaneously deform

Murrell and Gardel 2012

 

flow is driven by internal force imbalances

Passive

Active

Mayer et al. 2010

 

filament recycling: key to sustaining flows

F-actin filaments are highly dynamic

cellular mechanisms of filament recycling

disassembly takes places in 100 s

Kuey et al. JCB 2008

in vivo, the lifetime can be even shorter

during division

pre-division

Robin et al. 2012

Preventing filament recycling stalls flows or leads to cortical tearing

Jon Michaux

 

japlakinolide

goal: understand what sets flow rate

Passive

Active

stress

viscosity

approach: determine what sets network viscosity and internal stress

Probing passive viscosity of 2D filament networks in presence of applied force

frictional cross-link stress dissipation

measuring passive network deformation

position

x velocity

This led to typical 3 phase deformation

\tau_c
τc\tau_c
1/\eta
1/η1/\eta
G_0
G0G_0

G0 agrees with prior theoretical prediction

Head Levine Mackintosh, PRL 2003

estimate of viscosity of networks with cross-link relaxation

\tau_c
τc\tau_c
1/\eta
1/η1/\eta

what happens when we introduce highly simplified filament recycling?

filament recycling led to faster strains

position

x velocity

x velocity

no recycling

tau =  10s

Only with fast enough recycling, the network's effective viscosity decreased

\tau_x \approx 100\tau_c
τx100τc\tau_x \approx 100\tau_c

Measuring deformation and stress buildup in 2D active filament networks

build a minimal model of active networks

asymmetric filament compliance

L

Storm and Nelson PRE 2003

Active networks transiently contract

free boundary

and generate a transient net stress

fixed boundary

\tau_a
τa\tau_a

recycling tunes the steady state stress

\tau_a
τa\tau_a

biphasic dependence on recycling time

Combining passive viscosity and active stress to predict 1D flows

recap: estimates for stress and viscosity

recall the dependence on these two important timescales

equations for viscosity and stress can be combined to predict flow speed

\dot{\gamma}=\frac{\sigma}{\eta}
γ˙=ση\dot{\gamma}=\frac{\sigma}{\eta}

strain rate

viscosity

stress

to test our theory we generated simulations of polarized networks

x velocity

x velocity

position

1000s

\tau_r
τr\tau_r

33s

active region

charting steady state flow dependence on recycling timescale

dependence of flow speed on recycling qualitatively matches theoretical prediction

flow simulations recapitulated the predicted interplay of these two effects

inert
tearing

conclusions and future directions

this work offers a theoretical framework for understanding cortical flows

it sets up some experiments to test both  quantitative and qualitative predictions

it might help explain what needs to be controlled to generate synthetic systems

a theoretical framework

is "X experiment" important for tuning actomyosin flow?

experimental tests

synthetic systems

theseus old

By wmcfadd2

theseus old

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