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Long-range nematic order & anomalous fluctuations in collective - - PowerPoint PPT Presentation

Daiki Nishiguchi 1 Department of Physics, The University of Tokyo PhD student (D3) at Sano Lab. Ken H. Nagai (JAIST) Hugues Chat (CEA-Saclay, CSRC-Beijing) Masaki Sano (The University of Tokyo) Big Waves of Theoretical Sciences in Okinawa @


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Long-range nematic order & anomalous fluctuations in collective motion of filamentous bacteria

Daiki Nishiguchi

1

Department of Physics, The University of Tokyo PhD student (D3) at Sano Lab. Ken H. Nagai (JAIST) Hugues Chaté (CEA-Saclay, CSRC-Beijing) Masaki Sano (The University of Tokyo) Big Waves of Theoretical Sciences in Okinawa @ OIST

(arXiv:1604.04247)

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SLIDE 2

Collective motion of self-propelled particles

・Consume some kind of energy at the particle level (intrinsically nonequilibrium) ・Direction of motion is determined not solely by external force, but by internal degree of freedom.

2

■Examples of “Collective motion”

  • H. P. Zhang (2010)

school of jack tuna

Bacillus subtilis (枯草菌)

(www.youtube.com/watch?v=D6HdoIsLMFg)

bird flock

(www.youtube.com/watch?v=_tEFRAI9WSE)

polarity・shape・cell cycle... ■Characteristics of “self-propelled particles”

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SLIDE 3

A big wave in theoretical active matter

3 Vicsek, et al. Phys. Rev. Lett. 75, 6 (1995)

■Our motivation

↑average angle of particles inside radius R

: white noise uniformly distributed on density ↗

  • r

noise ↘

■Vicsek model

・Many studies on Vicsek-like models regarding collective behavior. ・However, no corresponding experiments yet. →We want to realize such experiments, if possible, with biological systems!

  • rder-disorder

transition (1st order)

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SLIDE 4

A big wave in theoretical active matter

3 Vicsek, et al. Phys. Rev. Lett. 75, 6 (1995)

■Our motivation

↑average angle of particles inside radius R

: white noise uniformly distributed on density ↗

  • r

noise ↘

■Vicsek model

・Many studies on Vicsek-like models regarding collective behavior. ・However, no corresponding experiments yet. →We want to realize such experiments, if possible, with biological systems!

  • rder-disorder

transition (1st order)

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SLIDE 5

A big wave in theoretical active matter

3 Vicsek, et al. Phys. Rev. Lett. 75, 6 (1995)

■Our motivation

↑average angle of particles inside radius R

: white noise uniformly distributed on density ↗

  • r

noise ↘

■Vicsek model

・Many studies on Vicsek-like models regarding collective behavior. ・However, no corresponding experiments yet. →We want to realize such experiments, if possible, with biological systems!

  • rder-disorder

transition (1st order)

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SLIDE 6

A big wave in theoretical active matter

3 Vicsek, et al. Phys. Rev. Lett. 75, 6 (1995)

■Our motivation

↑average angle of particles inside radius R

: white noise uniformly distributed on density ↗

  • r

noise ↘

■Vicsek model

・Many studies on Vicsek-like models regarding collective behavior. ・However, no corresponding experiments yet. →We want to realize such experiments, if possible, with biological systems!

  • rder-disorder

transition (1st order)

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SLIDE 7

4

■What is number fluctuation?

:# of observed particles at time t.

Properties: Giant Number Fluctuations (GNF)

:mean value :fluctuation (standard deviation) in equilibrium or random systems (Central limit theorem)

■What about in Vicsek model?

Chaté, et al. Eur. Phys. J. B 64, 451 (2008), Toner and Tu Phys. Rev. E 58, 4 (1998)

・Exponent larger than 0.5 in the homogeneous ordered state. →giant number fluctuation (GNF) ・Due to numerics and continuum theory:

Normal fluctuations

・Note: GNF is associated with Nambu- Goldstone modes in the ordered state.

Giant fluctuations

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SLIDE 8

Vicsek-like models with different symmetry

5

Vicsek'model ac-ve'nema-cs self1propelled'rods mo-lity polar apolar polar interac-on polar nema-c nema-c GNF'(numerics) ~'0.8 ~'0.8 ~'0.75 GNF'(con-nuum) 0.8'(exact) 1'(linear'theory) controversial

  • rder

True'LRO Quasi1LRO True1LRO

■Active nematics

e.g.:shaken rods Narayan, et al. Science, 317, 6 (2007).

cf:Vicsek model

Ngo, et al.

  • Phys. Rev. Lett. 113, 038302 (2014)

■Self-propelled rods nematic ordered state→

Ginelli, et al.

  • Phys. Rev. Lett. 104, 184502 (2010)

e.g.: Bacteria

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SLIDE 9

Exponents in continuum theories

6

■Vicsek model (Toner-Tu theory) ■Active nematics (Ramaswamy, et al)

・From symmetry arguments, hydrodynamic equations can be written: ・Dynamical renomalization group method can derive the exponent in 2D. ・Exponents of GNF is only known for linear theory. ・Some theoreticians believes that phenomenologies on active nematics and self-propelled rods should be the same.

Toner & Tu, PRL, 75, 23 (1995). Ramaswamy, et al. EPL, 62(2), 196, (2003). Mishra, et al., J. Stat. Mech., (2010) P02003. Vicsek'model ac-ve'nema-cs self1propelled'rods mo-lity polar apolar polar interac-on polar nema-c nema-c GNF'(numerics) ~'0.8 ~'0.8 ~'0.75 GNF'(con-nuum) 0.8'(exact) 1'(linear'theory) controversial

  • rder

True'LRO Quasi1LRO True1LRO ↑Symmetric Traceless part

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SLIDE 10

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Bacillus subtilis Zhang, Beér, Florin & Swinney (PNAS 2010) shaken rods Narayan, Ramaswamy & Menon (Science 2007)

These GNF is trivial because of their obvious structures.

Observation of “GNF”

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SLIDE 11

8

motility assay shaken polar disks

Schaller, et al. Nature, 467, 73 (2008) Schaller and Bausch, PNAS, 110, 4488 (2013) Deseigne, et al. Phys. Rev. Lett. 05,098001 (2010)

Observation of “GNF”

: variance

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SLIDE 12

motility assay shaken polar disks

Observed GNF in previous experiments

・Many experiments report GNF, although it’s not their main claims. ・However, these GNFs came from effects of boundary or clustering. →Looking at fluctuations not in homogeneous ordered states.

Bacillus subtilis shaken rods not ordered clustering not ordered boundary effects clustering banding clustering boundary effects Zhang, et al. PNAS, 107, 31, 13626 (2010).

Schaller and Bausch, PNAS, 110, 4488 (2013) Deseigne, et al.

  • Phys. Rev. Lett.

105, 098001 (2010)

Narayan, et al. Science, 317, 6 (2007).

→Realize experimental systems that exhibit homogeneous ordered states!

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SLIDE 13

What kind of experiments is required?

10

・As simple as possible to compare with numerical models. (avoid complicated interactions) ・Far from boundaries, a large number of particles to see statistics. →µm-scale particles are preferable. →microswimmers (e.g. self-propelled colloids or bacteria) (→Demonstrations with biological systems is important!) →Use bacteria. ・Flow fields of bacteria (pusher-type swimmer) destabilize ordered states. →Sandwich them between two walls to suppress flow. (The thinner, the more collisions. 2D is easier to observe.) ・To realize an ordered state, stronger alignment is required. →use higher-aspect-ratio particles ■Design principles for homogeneous ordered phases

Janus colloids

DN & M. Sano,

  • Phys. Rev. E

92, 052309 (2015)

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SLIDE 14

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particles:filamentous E. coli

High-aspect-ratio bacteria

10μm real speed

20-100 μm

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SLIDE 15

Experimental procedure

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Making filamentous cells Concentrate the suspension Confine in an observation device Observe and analyze

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SLIDE 16

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■procedure

・Add an antibiotic that inhibits cell division to the suspension of E.coli (strain: RP437 or RP4979). ・The length of E.coli can be controlled by changing the incubation time. real speed

Making filamentous cells

usual E.coli(2-3 µm) filamentous cells(30-40 µm) antibiotic

20 µm

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SLIDE 17

Experimental procedure

14

Making filamentous cells Concentrate the suspension Confine in an observation device Observe and analyze

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SLIDE 18

Experimental procedure

15

Making filamentous cells Concentrate the suspension Confine in an observation device Observe and analyze

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SLIDE 19

What kind of experiments is required?

16

・As simple as possible to compare with numerical models. (avoid complicated interactions) ・Far from boundaries, a large number of particles to see statistics. →µm-scale particles are preferable. →microswimmers (e.g. self-propelled colloids or bacteria) (→Demonstrations with biological systems is important!) →First introduce bacterial experiments!! ・Flow fields of bacteria (pusher-type swimmer) destabilize ordered states. →Sandwich them between two walls to suppress flow. (The thinner, the more collisions. 2D is easier to observe.) ・To realize an ordered state, stronger alignment is required. →use higher-aspect-ratio particles ■Experimental Systems for realizing homogeneous ordered states

Janus colloids

Nishiguchi & Sano,

  • Phys. Rev. E

92, 052309 (2015)

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SLIDE 20

Observation Device

17 Concentrated suspension Coverslip PDMS seal Objective lens excitation light

O2

・Oxygen is required for high motility. ・PDMS(polymeric organo silicon) is transparent to oxygen. ・Make patterns on PDMS so that excessive suspension escapes into halls. ・Gap between glass and PDMS: smaller than 2µm

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SLIDE 21

Experimental procedure

18

Making filamentous cells Concentrate the suspension Confine in an observation device Observe and analyze

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SLIDE 22

Swimming at low density

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・Used strain: RP4979(ΔcheY, YFP), chemotactic mutant →no tumbling, swim smoothly, exclude artifacts.

some cells swim together after collisions→

50μm

x40 objective lens captured@10fps, real speed wildtype with tumbling mutants without tumbling →motility is polar!

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SLIDE 23

Interactions are nematic!

20

time 0 s 0.2 s 0.4 s 0.6 s

20µm

acute angle →parallel alignment

  • btuse angle

→anti-parallel alignment

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SLIDE 24

Ordered state at high density

21

・Each particle has polarity. ・Collisions lead to global nematic order. →self-propelled rods model ! real speed captured@10fps

Vicsek'model ac-ve'nema-cs self1propelled'rods mo-lity polar apolar polar interac-on polar nema-c nema-c exponent'of'GNF 0.8 ~'0.8 ~'0.75 100 µm

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SLIDE 25

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Observed with lower magnification

Ordered state in large area! →measure GNF!! x3 fast x10 objective captured@10fps

100μm

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SLIDE 26

Disordered state for comparison

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x3 fast x10 objective captured @10fps

100μm

Disordered state is random. Number fluctuation obeys (central limit theorem)

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SLIDE 27

Results of GNF measurement

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■Experimental Result

Ginelli, et al. Phys. Rev. Lett. 104, 184502 (2010)

・n: number of pixels in a box that bacteria exist (1 bacterium ~ 100 pixels). ・We obtained giant number fluctuations over 3-4 decades.

→fitting: 0.632±0.014

■Self-propelled rods model

(small due to exclusive volume?)

10

1

10

2

10

3

10

4

10

5

10

6

5 10 20 30

  • rdered

disordered →fitting: 0.511±0.012

Exponent ~ 0.75

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SLIDE 28

What kind of “order” in ordered states?

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■Do they have long-range order? ■How to distinguish

・Plot nematic order parameter against calculation box size and see how they decay. →quasi long-range order: algebraic decay (with small exponent) toward 0 true long-range order: converge to a positive value

LRO:'Long1Range'Order

Vicsek'model ac-ve'nema-cs self1propelled'rods mo-lity polar apolar polar interac-on polar nema-c nema-c GNF'(numerics) ~'0.8 ~'0.8 ~'0.75 GNF'(con-nuum) 0.8'(exact) 1'(linear'theory) controversial

  • rder

True'LRO Quasi1LRO True1LRO

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SLIDE 29

Long-range order?

26

evaluate

■How to distinguish

・Plot nematic order parameter against calculation box size and see how they decay. →quasi long-range order: algebraic decay (with small exponent) toward 0 true long-range order: converge to a positive value

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SLIDE 30

27

evaluate

Long-range order?

■How to distinguish

・Plot nematic order parameter against calculation box size and see how they decay. →quasi long-range order: algebraic decay (with small exponent) toward 0 true long-range order: converge to a positive value

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SLIDE 31

28

evaluate

Long-range order?

■How to distinguish

・Plot nematic order parameter against calculation box size and see how they decay. →quasi long-range order: algebraic decay (with small exponent) toward 0 true long-range order: converge to a positive value

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SLIDE 32

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■Evaluating nematic order ・Direct calculation of S requires detecting all the cells (difficult). ・Evaluate by ”structure tensor” method (find max/min of intensity gradient). :image data in the box e.g.: Coherency: →regard this as nematic order S →take average of Coherency

Rezakhaniha, et al.

  • Biomech. Model. Mechanobiol. 11, 3-4, 461 (2011)

Long-range order?

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SLIDE 33

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■Nematic Order vs Box Size ■Self-propelled rods model

Long-range order? → YES!!

Ginelli, et al.

  • Phys. Rev. Lett.

104, 184502 (2010)

・Slower decay than algebraic decay. ・Converge to a positive value. →true long-range order! (consistent with self-propelled rods model) ・Obtained GNF should remain in large system size limit.

10

3

10

4

10

5

10

  • 10
  • 10
  • S (m)

10

3

10

4

10

5

10

6

0.50 0.52 0.55

S (m2)

due to spatial inhomogeneity

  • f experimental setup
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SLIDE 34

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■Correlation function

Correlation of nematic director fluctuations

  • [µm]

[µm]

10 100 800 0.03 0.10 1.00

Distance [µm] Correlatioin

  • 0.3

Correlation should reflect the properties of Nambu-Goldstone mode. →What is the theoretical dependence of Corr(r)? (on-going) power-law? local director↑ ↑global(mean) director fluctua-on'of'director:

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SLIDE 35

Further experimental tests on Vicsek Physics

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Janus'par-cles filamentous'bacteria ■Classification of Vicsek-like models (Poster'by'J.'Iwasawa) (This'Talk'&'my'poster)

Vicsek'model ac-ve'nema-cs self1propelled'rods mo-lity polar apolar polar interac-on polar nema-c nema-c GNF'(numerics) ~'0.8 ~'0.8 ~'0.75 GNF'(con-nuum) 0.8'(exact) 1'(linear'theory) controversial

  • rder

True'LRO Quasi1LRO True1LRO Janus colloids

DN & M. Sano,

  • Phys. Rev. E

92, 052309 (2015)

  • E
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SLIDE 36

Summary on filamentous bacteria

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Summary

・We have realized an ordered state of collective motion with a biological system using filamentous cells of E. coli. ・The obtained ordered state has true long-range order. ・“GNF in the sense of numerics/theories”(not from clustering etc.), which had not been experimentally observed, is observed over 3-4 decades. ・All the results are consistent with self-propelled rods model (numerics).

32

Prospects

・See the phase transition. ・Other properties suggested by numerics & theory (superdiffusion, dispersion, etc.) ・See how aspect ratio affects collective dynamics.

Acknowledgement

・Transformation of E.coli (fluorescent protein): Yusuke Maeda (Kyushu Univ.)

Vicsek'model ac-ve'nema-cs self1propelled'rods mo-lity polar apolar polar interac-on polar nema-c nema-c GNF'(numerics) ~'0.8 ~'0.8 ~'0.75 GNF'(con-nuum) 0.8'(exact) 1'(linear'theory) controversial

  • rder

True'LRO Quasi1LRO True1LRO

Preprint available!! → arXiv:1604.04247