SlideShare a Scribd company logo
Stochastic modelling of zebrafish locomotion :
collective motion from the bottom-up
Adam Zienkiewicz1 • David A. W. Barton1 • Maurizio Porfiri2 • Mario di Bernardo1,3
1Department of Engineering Mathematics, University of Bristol, UK
2Department of Mechanical and Aerospace Engineering,
New York University Polytechnic School of Engineering, USA
3Department of Information and Computer Engineering,
University of Naples Federico II, Italy
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
Overview
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(1 / 25)
Background and motivation Why study animal motion? Why (zebra)fish?
Individual model
…motion of (isolated) zebrafish
Multi-agent modeling
…modeling schools and collective motion
Emergence of leadership in groups
…towards methods of control
• requirements of a new model
• experiments, observations
• modeling and results
• observations and experiments
• model extension & results
• macro-level dynamics
Data-driven stochastic modeling of
zebrafish locomotion.
Zienkiewicz et al. , J. Math. Biology (2014)
Introduction
• Recent stochastic models of fish locomotion
- “Persistent Turning Walker” (PTW) model : Kulia mugil
displacement described in terms of turning speed and its autocorrelation
- later extended to multiple fish, modelling collective behaviour (same authors)
...motivation and background
Gautrais et al. (2009) J. Math. Biol.
58(3) and (2012) PLoS Comp. Biol. 8(9)
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(2 / 25)
Berdahl et al. (2013). Science, 339(6119)
• Speed regulation as a primary response mechanism
- decomposition of interaction forces within groups of small, shoaling fish (golden shiners)
Katz et al. and Herbert-Read et al. (2011) PNAS 108(46)
- emergent sensing (distributed / decentralised) of environment via speed regulation
Berdahl et al. (2013) Science, 339(6119)
Katz et al. (2011) PNAS 108(46)
Model species ...zebrafish
Zebrafish (Danio rerio) - Increasingly predominant species for neurobiological,
developmental and behavioural laboratory studies
small ● fast reproductive cycle ● genetic homology
high stocking density ● strong social groups
≈ 3 cm
© Azul 2005
cf. Kulia Mugil ≈ 20-25 cm
No suitable model! - Can similar stochastic PTW type model be used?
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(3 / 25)
Introduction ...objectives
Using data collected from video tracking of individual trajectories,
develop a data-driven model framework describing the motion
characteristics of individual zebrafish
• Adapt Gautrais’ PTW (constant speed) model of fish locomotion to capture
salient swimming features of small, shoaling fish with burst-and-coast
swimming mode
- augment stochastic differential model with additional dynamic speed process
- identify sufficient key metrics required to describe fish locomotion
- allow for and characterise interactions with environment (wall-avoidance)
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(4 / 25)
• Model framework features to provide foundation for extension to multiple
fish dynamics : biologically realistic models of (zebrafish) collective motion
- speed regulation : dynamic speed interaction models
vs. (canonical) constant speed
Experiments
Zebrafish trajectory data -
...data capture : automated tracking
• 10 isolated individuals observed for 5 min. Each
- shallow (10 cm), square tank (120 x 120 x20 cm)
• overhead video capture - 30 fps  5 fps (Hz)
• automated visual tracking of centroid
(point) position of fish in tank
(Kalman filter)
(rate of change of heading angle of vt )
VIDEO: automated position
tracking of live zebrafish
Automated fish (particle) tracking
• compute velocity
• speed
• turning speed
Dynamical Systems Laboratory (NYU)
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(5 / 25)
Observations ...zebrafish trajectories
Raw trajectory data: 10 zebrafish, 5 min observations Speed (ut)
Individual zebrafish exhibit a variety of locomotory patterns:
- smooth, fluid turning
- erratic (stop / start) with sharp or spiralling turning
- wall following (thigmotaxis)
BL.s-1
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(6 / 25)
Observations ...segment trajectories
Swimming segment trajectories (coloured by parent fish ID F1...F10)
‘Swimming’ data isolated from individual zebrafish : 28 segments of equal duration
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(7 / 25)
Trajectory analysis ...primary characteristics
Time series, distribution and autocorrelation (segment S9)
Time lag (s)
ut and ωt characterised by their distributions and autocorrelation (ACF)
- approximate with normal distributions and
Typically sharper
than Gaussian
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(8 / 25)
Trajectory analysis ...primary characteristics
Speed / Turning speed cross correlation (log-frequency : all segments)
Strong correlation within ut and ωt joint distribution across all segments
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(9 / 25)
Individual Model ...stochastic differential model
Capture salient characteristics of trajectory data using mean-reverting
stochastic processes : coupled stochastic differential equations
Wiener processes: dWt and dZt
Speed:
Ornstein-Uhlenbeck (type) processes:
Couple equations
with function:
Wall interactions
via bias function:
• mean speed: , mean turning speed:
• exp. decaying ACF with rates:
• process volatilities: and
6 variable parameters :
estimated from each
swimming segment
Turning-speed:
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(10 / 25)
Individual Model (2) ...wall avoidance and SDE coupling
Boundary effect on turning speed (S27)Quantify effect of wall interactions
on trajectories:
- induced fwd. acceleration : inconclusive
- induced turning : trajectories bent away from
wall dependent on projected collision angle
Define a coupling function to restrict the volatility of Ωt process
as a function of Ut
• fc → σ0 as Ut → 0 (upper bounded)
• fc → 0 as Ut → ∞ (lower bounded)
• fc → σω /2 as Ut → μu (σω estimated from data)
estimate constants (A,B σ0 )
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(11 / 25)
Results ...segment-wise calibration and simulation
Qualitative comparisons : S13 vs. RW13
Speed Turning speed
Dist.ACF
Calibrate SDE parameters [μ,σ,ϑ]u,ω from experimental speed / turning speed data
max. likelihood est.
assume standard
(Gaussian) O-U processes
dU, dΩ numerically integrated
(Euler-Maruyama)
 generate random walks
Trajectories : S13 vs. RW
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(12 / 25)
Results ...process coupling
Coupling function fc recovers joint distribution of Ut and Ωt
Speed / Turning speed cross correlation (composite of all segments )
experimental data simulated random walkers
...upper volatility bound σ0 fixed, μu and σω vary between RW segments
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(13 / 25)
Results ...segment simulation
Example simulation (2x real-time)
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(14 / 25)
Results ...calibrating individual fish
Trajectory comparison (individual fish) insufficient swimming data for F4 & F8
Average segment parameters used for calibration of (8) individuals
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(15 / 25)
Modelling a shoal ... multi-agent models of collective behaviour
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(16 / 25)
Pairs of zebrafish swimming in circular shallow tank
(45 cm radius, 10cm water depth)
Experiments:
• 18 observations of unique zebrafish pairs
• 20 min observations (30Hz sample freq.)
• automated tracking + manual repair
• samples proximal (< 2 BL) to walls omitted
pair
What kind of behaviour (rules) can we infer from observations of fish swimming together?
Attraction? Repulsion? Alignment?
Analysis of trajectory data can reveal
pairwise interactions in terms of
‘social forces’ (accelerations)
Jun – Aug 2014
Hold up! …start with two fish
(6 hrs)
Inferring interaction behaviour ... social forces
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(17 / 25)
Katz et al. (2011) PNAS 108(46)Golden shiners (14 x 56 min @ 30Hz)
Zebrafish (2 x 20 min @ 30Hz)
5 cm
(juvenile)
3 cm
(adult)
Zienkiewicz et al. (incomplete data)
Inferring interaction behaviour ... social forces
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(18 / 25)
Golden shiners (14 x 56 min @ 30Hz)
Zebrafish (2 x 20 min @ 30Hz)
5 cm
(juvenile)
3 cm
(adult)
Golden shiners
Zebrafish
Alignment :
an emergent phenomena?
Data suggests turning is dependent
only on relative position
….not orientation
 no explicit alignment ‘rule’
Interaction model ...toy model of speed / turning forces
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(19 / 25)
Interpolate forces in primary response direction: Model potential:
Angular forceTangential (speed) force
Interaction model (2) ...conclusions
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(20 / 25)
Speed:
Turning speed:
Modified SDEs:
Acceleration (force) due to pair-wise interactions:
(Similarly for angular acc. )
Interaction network adjacency matrix:
Voronoi neighbourhood
(Voronoi neighbourhood, radial proximity
networks, estimated visual networks)
Multi-agent simulations ...2-fish example
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(21 / 25)
Global observables ...2-fish example
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(22 / 25)
Polarisation (P)
Milling (M)
Cohesion (P)
Mean nearest-
neighbour
distance (MNND)
Live zebrafish Simulation
2 fish, 20 mins @ 30fps 2 agents, 20 mins @ 30 Hz
A set of measures / order parameters which describe the global, or macro-scale dynamics
Relative alignment
Rotation around a
common centre of mass
Rotation around a
common centre of mass
Multi-agent simulations ... 1000 fish example !!
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(23 / 25)
Leadership and collective decision making
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(24 / 25)
How can collective dynamics
be modulated by the
presence of a subset of
Informed individuals?
Emergent leadership in the
absence of explicit signals....
• zero turning (translation)
• constant turning (circular)
• ‘blind’ agent
• agent(s) with preferential
heading direction
Practical examples:
foraging, migration,
danger awareness
...artificial control ?
Conclusions
• direct calibration from experimental data, inc. boundary avoidance
- produce simulated trajectories with comparable curvature
- capture ‘passive’ wall following behaviour with ϑω /σω dependence
• describe zebrafish locomotion with an extended PTW model
- characterised by autoregressive, stochastic processes for
both speed and turning speed
- stochastic speed process more suitable for burst-and-coast
swimming mode of small, schooling fish
• model framework allows explicit inclusion of both speed and
turning speed modulation as responses to dynamic environment
- equilibrium bias of both speed and turning speed can be evolved
to simulate linear accelerations and torques (independently)
- infer interaction ‘forces’ to model group behaviour
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014
(25 / 25)
Acknowledgements
My supervisors:
Mario di Bernardo
David Barton
Maurizio Porfiri
My sponsors (U.K.):
Dynamical Systems Laboratory:
(New York University Polytechnic School of Engineering)
Sachit Butail
Fabrizio Ladu
...thank you for listening
Adam Zienkiewicz Stochastic modelling of zebrafish locomotion
Complexity (BCCS) Seminar – 16 December 2014

More Related Content

Similar to BCCS_seminar_Dec2014

Autonomous Underwater Vehicles - Copy (3).pptx
Autonomous Underwater Vehicles - Copy (3).pptxAutonomous Underwater Vehicles - Copy (3).pptx
Autonomous Underwater Vehicles - Copy (3).pptx
animeshmahatajgm2001
 
Emerging 3D Display Technologies
Emerging 3D Display TechnologiesEmerging 3D Display Technologies
Emerging 3D Display Technologies
Matt Hirsch - MIT Media Lab
 
Hydrodynamics of Dolphin Caudal Fins
Hydrodynamics of Dolphin Caudal FinsHydrodynamics of Dolphin Caudal Fins
Hydrodynamics of Dolphin Caudal FinsEric Zacharia
 
Biohybrid Robotic Jellyfish for Future Applications in Ocean Monitoring
Biohybrid Robotic Jellyfish for Future Applications in Ocean MonitoringBiohybrid Robotic Jellyfish for Future Applications in Ocean Monitoring
Biohybrid Robotic Jellyfish for Future Applications in Ocean Monitoring
inside-BigData.com
 
(May 9, 2024) Enhanced Ultrafast Vector Flow Imaging (VFI) Using Multi-Angle ...
(May 9, 2024) Enhanced Ultrafast Vector Flow Imaging (VFI) Using Multi-Angle ...(May 9, 2024) Enhanced Ultrafast Vector Flow Imaging (VFI) Using Multi-Angle ...
(May 9, 2024) Enhanced Ultrafast Vector Flow Imaging (VFI) Using Multi-Angle ...
Scintica Instrumentation
 
Seminar at University of Graz (12.05.2015)
Seminar at University of Graz (12.05.2015)Seminar at University of Graz (12.05.2015)
Seminar at University of Graz (12.05.2015)
Alexey Isavnin
 
HHMI Day 2016 Poster_final
HHMI Day 2016 Poster_finalHHMI Day 2016 Poster_final
HHMI Day 2016 Poster_finalFrancis Lin
 
Automated sorting of parasitic worms using sine-wave channels
Automated sorting of parasitic worms using sine-wave channelsAutomated sorting of parasitic worms using sine-wave channels
Automated sorting of parasitic worms using sine-wave channels
Iowa State University
 
Basu_CV
Basu_CVBasu_CV
Basu_CV
Saikat Basu
 
20140616 19 depestele-physical_impact_vfinalpresented
20140616 19 depestele-physical_impact_vfinalpresented20140616 19 depestele-physical_impact_vfinalpresented
20140616 19 depestele-physical_impact_vfinalpresented
Jochen Depestele
 
TeZ - SPECTRAL SENSORIUM - ART/SCIENCE RESEARCH 2010-2015
TeZ - SPECTRAL SENSORIUM - ART/SCIENCE RESEARCH 2010-2015TeZ - SPECTRAL SENSORIUM - ART/SCIENCE RESEARCH 2010-2015
TeZ - SPECTRAL SENSORIUM - ART/SCIENCE RESEARCH 2010-2015
TeZ Martinucci
 
Fisher Kernel based Relevance Feedback for Multimodal Video Retrieval
Fisher Kernel based Relevance Feedback for Multimodal Video RetrievalFisher Kernel based Relevance Feedback for Multimodal Video Retrieval
Fisher Kernel based Relevance Feedback for Multimodal Video Retrieval
Ionut Mironica
 
Vessels delineation in retinal 
images using COSFIRE filters
Vessels delineation in retinal 
images using COSFIRE filtersVessels delineation in retinal 
images using COSFIRE filters
Vessels delineation in retinal 
images using COSFIRE filters
Nicola Strisciuglio
 
ICSU World Data System for scientific research
ICSU World Data System for scientific researchICSU World Data System for scientific research
ICSU World Data System for scientific research
SSA KPI
 
Whispers of Speckles ( Part I: Building Computational Imaging Frameworks for ...
Whispers of Speckles (Part I: Building Computational Imaging Frameworks for ...Whispers of Speckles (Part I: Building Computational Imaging Frameworks for ...
Whispers of Speckles ( Part I: Building Computational Imaging Frameworks for ...
Debdoot Sheet
 
Automatic reading cr39
Automatic reading cr39Automatic reading cr39
Automatic reading cr39
MOAYYAD ALSSABBAGH
 
Arrows presentation-emra-2015
Arrows presentation-emra-2015Arrows presentation-emra-2015
Arrows presentation-emra-2015
Benedetto Allotta
 
Acoustic Modal Analaysis Hydrofoils.pdf
Acoustic Modal Analaysis Hydrofoils.pdfAcoustic Modal Analaysis Hydrofoils.pdf
Acoustic Modal Analaysis Hydrofoils.pdf
SRIJNA SINGH
 
Presentation CIE619
Presentation CIE619 Presentation CIE619
Presentation CIE619
Sharath Chandra
 

Similar to BCCS_seminar_Dec2014 (20)

Autonomous Underwater Vehicles - Copy (3).pptx
Autonomous Underwater Vehicles - Copy (3).pptxAutonomous Underwater Vehicles - Copy (3).pptx
Autonomous Underwater Vehicles - Copy (3).pptx
 
Emerging 3D Display Technologies
Emerging 3D Display TechnologiesEmerging 3D Display Technologies
Emerging 3D Display Technologies
 
Thesis_de_Meulenaer
Thesis_de_MeulenaerThesis_de_Meulenaer
Thesis_de_Meulenaer
 
Hydrodynamics of Dolphin Caudal Fins
Hydrodynamics of Dolphin Caudal FinsHydrodynamics of Dolphin Caudal Fins
Hydrodynamics of Dolphin Caudal Fins
 
Biohybrid Robotic Jellyfish for Future Applications in Ocean Monitoring
Biohybrid Robotic Jellyfish for Future Applications in Ocean MonitoringBiohybrid Robotic Jellyfish for Future Applications in Ocean Monitoring
Biohybrid Robotic Jellyfish for Future Applications in Ocean Monitoring
 
(May 9, 2024) Enhanced Ultrafast Vector Flow Imaging (VFI) Using Multi-Angle ...
(May 9, 2024) Enhanced Ultrafast Vector Flow Imaging (VFI) Using Multi-Angle ...(May 9, 2024) Enhanced Ultrafast Vector Flow Imaging (VFI) Using Multi-Angle ...
(May 9, 2024) Enhanced Ultrafast Vector Flow Imaging (VFI) Using Multi-Angle ...
 
Seminar at University of Graz (12.05.2015)
Seminar at University of Graz (12.05.2015)Seminar at University of Graz (12.05.2015)
Seminar at University of Graz (12.05.2015)
 
HHMI Day 2016 Poster_final
HHMI Day 2016 Poster_finalHHMI Day 2016 Poster_final
HHMI Day 2016 Poster_final
 
Automated sorting of parasitic worms using sine-wave channels
Automated sorting of parasitic worms using sine-wave channelsAutomated sorting of parasitic worms using sine-wave channels
Automated sorting of parasitic worms using sine-wave channels
 
Basu_CV
Basu_CVBasu_CV
Basu_CV
 
20140616 19 depestele-physical_impact_vfinalpresented
20140616 19 depestele-physical_impact_vfinalpresented20140616 19 depestele-physical_impact_vfinalpresented
20140616 19 depestele-physical_impact_vfinalpresented
 
TeZ - SPECTRAL SENSORIUM - ART/SCIENCE RESEARCH 2010-2015
TeZ - SPECTRAL SENSORIUM - ART/SCIENCE RESEARCH 2010-2015TeZ - SPECTRAL SENSORIUM - ART/SCIENCE RESEARCH 2010-2015
TeZ - SPECTRAL SENSORIUM - ART/SCIENCE RESEARCH 2010-2015
 
Fisher Kernel based Relevance Feedback for Multimodal Video Retrieval
Fisher Kernel based Relevance Feedback for Multimodal Video RetrievalFisher Kernel based Relevance Feedback for Multimodal Video Retrieval
Fisher Kernel based Relevance Feedback for Multimodal Video Retrieval
 
Vessels delineation in retinal 
images using COSFIRE filters
Vessels delineation in retinal 
images using COSFIRE filtersVessels delineation in retinal 
images using COSFIRE filters
Vessels delineation in retinal 
images using COSFIRE filters
 
ICSU World Data System for scientific research
ICSU World Data System for scientific researchICSU World Data System for scientific research
ICSU World Data System for scientific research
 
Whispers of Speckles ( Part I: Building Computational Imaging Frameworks for ...
Whispers of Speckles (Part I: Building Computational Imaging Frameworks for ...Whispers of Speckles (Part I: Building Computational Imaging Frameworks for ...
Whispers of Speckles ( Part I: Building Computational Imaging Frameworks for ...
 
Automatic reading cr39
Automatic reading cr39Automatic reading cr39
Automatic reading cr39
 
Arrows presentation-emra-2015
Arrows presentation-emra-2015Arrows presentation-emra-2015
Arrows presentation-emra-2015
 
Acoustic Modal Analaysis Hydrofoils.pdf
Acoustic Modal Analaysis Hydrofoils.pdfAcoustic Modal Analaysis Hydrofoils.pdf
Acoustic Modal Analaysis Hydrofoils.pdf
 
Presentation CIE619
Presentation CIE619 Presentation CIE619
Presentation CIE619
 

BCCS_seminar_Dec2014

  • 1. Stochastic modelling of zebrafish locomotion : collective motion from the bottom-up Adam Zienkiewicz1 • David A. W. Barton1 • Maurizio Porfiri2 • Mario di Bernardo1,3 1Department of Engineering Mathematics, University of Bristol, UK 2Department of Mechanical and Aerospace Engineering, New York University Polytechnic School of Engineering, USA 3Department of Information and Computer Engineering, University of Naples Federico II, Italy Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014
  • 2. Overview Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (1 / 25) Background and motivation Why study animal motion? Why (zebra)fish? Individual model …motion of (isolated) zebrafish Multi-agent modeling …modeling schools and collective motion Emergence of leadership in groups …towards methods of control • requirements of a new model • experiments, observations • modeling and results • observations and experiments • model extension & results • macro-level dynamics Data-driven stochastic modeling of zebrafish locomotion. Zienkiewicz et al. , J. Math. Biology (2014)
  • 3. Introduction • Recent stochastic models of fish locomotion - “Persistent Turning Walker” (PTW) model : Kulia mugil displacement described in terms of turning speed and its autocorrelation - later extended to multiple fish, modelling collective behaviour (same authors) ...motivation and background Gautrais et al. (2009) J. Math. Biol. 58(3) and (2012) PLoS Comp. Biol. 8(9) Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (2 / 25) Berdahl et al. (2013). Science, 339(6119) • Speed regulation as a primary response mechanism - decomposition of interaction forces within groups of small, shoaling fish (golden shiners) Katz et al. and Herbert-Read et al. (2011) PNAS 108(46) - emergent sensing (distributed / decentralised) of environment via speed regulation Berdahl et al. (2013) Science, 339(6119) Katz et al. (2011) PNAS 108(46)
  • 4. Model species ...zebrafish Zebrafish (Danio rerio) - Increasingly predominant species for neurobiological, developmental and behavioural laboratory studies small ● fast reproductive cycle ● genetic homology high stocking density ● strong social groups ≈ 3 cm © Azul 2005 cf. Kulia Mugil ≈ 20-25 cm No suitable model! - Can similar stochastic PTW type model be used? Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (3 / 25)
  • 5. Introduction ...objectives Using data collected from video tracking of individual trajectories, develop a data-driven model framework describing the motion characteristics of individual zebrafish • Adapt Gautrais’ PTW (constant speed) model of fish locomotion to capture salient swimming features of small, shoaling fish with burst-and-coast swimming mode - augment stochastic differential model with additional dynamic speed process - identify sufficient key metrics required to describe fish locomotion - allow for and characterise interactions with environment (wall-avoidance) Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (4 / 25) • Model framework features to provide foundation for extension to multiple fish dynamics : biologically realistic models of (zebrafish) collective motion - speed regulation : dynamic speed interaction models vs. (canonical) constant speed
  • 6. Experiments Zebrafish trajectory data - ...data capture : automated tracking • 10 isolated individuals observed for 5 min. Each - shallow (10 cm), square tank (120 x 120 x20 cm) • overhead video capture - 30 fps  5 fps (Hz) • automated visual tracking of centroid (point) position of fish in tank (Kalman filter) (rate of change of heading angle of vt ) VIDEO: automated position tracking of live zebrafish Automated fish (particle) tracking • compute velocity • speed • turning speed Dynamical Systems Laboratory (NYU) Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (5 / 25)
  • 7. Observations ...zebrafish trajectories Raw trajectory data: 10 zebrafish, 5 min observations Speed (ut) Individual zebrafish exhibit a variety of locomotory patterns: - smooth, fluid turning - erratic (stop / start) with sharp or spiralling turning - wall following (thigmotaxis) BL.s-1 Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (6 / 25)
  • 8. Observations ...segment trajectories Swimming segment trajectories (coloured by parent fish ID F1...F10) ‘Swimming’ data isolated from individual zebrafish : 28 segments of equal duration Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (7 / 25)
  • 9. Trajectory analysis ...primary characteristics Time series, distribution and autocorrelation (segment S9) Time lag (s) ut and ωt characterised by their distributions and autocorrelation (ACF) - approximate with normal distributions and Typically sharper than Gaussian Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (8 / 25)
  • 10. Trajectory analysis ...primary characteristics Speed / Turning speed cross correlation (log-frequency : all segments) Strong correlation within ut and ωt joint distribution across all segments Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (9 / 25)
  • 11. Individual Model ...stochastic differential model Capture salient characteristics of trajectory data using mean-reverting stochastic processes : coupled stochastic differential equations Wiener processes: dWt and dZt Speed: Ornstein-Uhlenbeck (type) processes: Couple equations with function: Wall interactions via bias function: • mean speed: , mean turning speed: • exp. decaying ACF with rates: • process volatilities: and 6 variable parameters : estimated from each swimming segment Turning-speed: Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (10 / 25)
  • 12. Individual Model (2) ...wall avoidance and SDE coupling Boundary effect on turning speed (S27)Quantify effect of wall interactions on trajectories: - induced fwd. acceleration : inconclusive - induced turning : trajectories bent away from wall dependent on projected collision angle Define a coupling function to restrict the volatility of Ωt process as a function of Ut • fc → σ0 as Ut → 0 (upper bounded) • fc → 0 as Ut → ∞ (lower bounded) • fc → σω /2 as Ut → μu (σω estimated from data) estimate constants (A,B σ0 ) Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (11 / 25)
  • 13. Results ...segment-wise calibration and simulation Qualitative comparisons : S13 vs. RW13 Speed Turning speed Dist.ACF Calibrate SDE parameters [μ,σ,ϑ]u,ω from experimental speed / turning speed data max. likelihood est. assume standard (Gaussian) O-U processes dU, dΩ numerically integrated (Euler-Maruyama)  generate random walks Trajectories : S13 vs. RW Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (12 / 25)
  • 14. Results ...process coupling Coupling function fc recovers joint distribution of Ut and Ωt Speed / Turning speed cross correlation (composite of all segments ) experimental data simulated random walkers ...upper volatility bound σ0 fixed, μu and σω vary between RW segments Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (13 / 25)
  • 15. Results ...segment simulation Example simulation (2x real-time) Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (14 / 25)
  • 16. Results ...calibrating individual fish Trajectory comparison (individual fish) insufficient swimming data for F4 & F8 Average segment parameters used for calibration of (8) individuals Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (15 / 25)
  • 17. Modelling a shoal ... multi-agent models of collective behaviour Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (16 / 25) Pairs of zebrafish swimming in circular shallow tank (45 cm radius, 10cm water depth) Experiments: • 18 observations of unique zebrafish pairs • 20 min observations (30Hz sample freq.) • automated tracking + manual repair • samples proximal (< 2 BL) to walls omitted pair What kind of behaviour (rules) can we infer from observations of fish swimming together? Attraction? Repulsion? Alignment? Analysis of trajectory data can reveal pairwise interactions in terms of ‘social forces’ (accelerations) Jun – Aug 2014 Hold up! …start with two fish (6 hrs)
  • 18. Inferring interaction behaviour ... social forces Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (17 / 25) Katz et al. (2011) PNAS 108(46)Golden shiners (14 x 56 min @ 30Hz) Zebrafish (2 x 20 min @ 30Hz) 5 cm (juvenile) 3 cm (adult) Zienkiewicz et al. (incomplete data)
  • 19. Inferring interaction behaviour ... social forces Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (18 / 25) Golden shiners (14 x 56 min @ 30Hz) Zebrafish (2 x 20 min @ 30Hz) 5 cm (juvenile) 3 cm (adult) Golden shiners Zebrafish Alignment : an emergent phenomena? Data suggests turning is dependent only on relative position ….not orientation  no explicit alignment ‘rule’
  • 20. Interaction model ...toy model of speed / turning forces Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (19 / 25) Interpolate forces in primary response direction: Model potential: Angular forceTangential (speed) force
  • 21. Interaction model (2) ...conclusions Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (20 / 25) Speed: Turning speed: Modified SDEs: Acceleration (force) due to pair-wise interactions: (Similarly for angular acc. ) Interaction network adjacency matrix: Voronoi neighbourhood (Voronoi neighbourhood, radial proximity networks, estimated visual networks)
  • 22. Multi-agent simulations ...2-fish example Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (21 / 25)
  • 23. Global observables ...2-fish example Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (22 / 25) Polarisation (P) Milling (M) Cohesion (P) Mean nearest- neighbour distance (MNND) Live zebrafish Simulation 2 fish, 20 mins @ 30fps 2 agents, 20 mins @ 30 Hz A set of measures / order parameters which describe the global, or macro-scale dynamics Relative alignment Rotation around a common centre of mass Rotation around a common centre of mass
  • 24. Multi-agent simulations ... 1000 fish example !! Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (23 / 25)
  • 25. Leadership and collective decision making Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (24 / 25) How can collective dynamics be modulated by the presence of a subset of Informed individuals? Emergent leadership in the absence of explicit signals.... • zero turning (translation) • constant turning (circular) • ‘blind’ agent • agent(s) with preferential heading direction Practical examples: foraging, migration, danger awareness ...artificial control ?
  • 26. Conclusions • direct calibration from experimental data, inc. boundary avoidance - produce simulated trajectories with comparable curvature - capture ‘passive’ wall following behaviour with ϑω /σω dependence • describe zebrafish locomotion with an extended PTW model - characterised by autoregressive, stochastic processes for both speed and turning speed - stochastic speed process more suitable for burst-and-coast swimming mode of small, schooling fish • model framework allows explicit inclusion of both speed and turning speed modulation as responses to dynamic environment - equilibrium bias of both speed and turning speed can be evolved to simulate linear accelerations and torques (independently) - infer interaction ‘forces’ to model group behaviour Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014 (25 / 25)
  • 27. Acknowledgements My supervisors: Mario di Bernardo David Barton Maurizio Porfiri My sponsors (U.K.): Dynamical Systems Laboratory: (New York University Polytechnic School of Engineering) Sachit Butail Fabrizio Ladu ...thank you for listening Adam Zienkiewicz Stochastic modelling of zebrafish locomotion Complexity (BCCS) Seminar – 16 December 2014