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Compositionality - Compositionality - Combinations Combinations of Muscle Synergies in the of Muscle Synergies in the Construction of Motor Construction of Motor Behavior Behavior Emilio Bizzi Massachusetts Institute of Technology
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Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

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Page 1: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Compositionality - CombinationsCompositionality - Combinations

of Muscle Synergies in theof Muscle Synergies in the

Construction of Motor BehaviorConstruction of Motor Behavior

Emilio BizziMassachusetts Institute of Technology

Page 2: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Compositionality: The genetic code and language are examplesof systems in which discreteelements can generate a large number of meaningful entitiesthat are quite distinct from thoseof their elements

Page 3: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Modularity

• Does the vertebrate motor system construct movements combining discrete modular elements?

Page 4: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

The Structure of Skeletal Muscle

Page 5: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

EMG recordings from 16 leg muscles

Page 6: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

a

b

c

synergy

muscles

Evidence for muscle synergies ?

• If a group of muscles is controlled as a unit, i.e. as a synergy, then the level of activity of those muscles should be correlated

Page 7: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Extraction algorithm

• We developed an iterative algorithm to extract a set of time-varying synergies that minimize the total reconstruction error

[d’Avella & Tresch, NIPS 14]

Page 8: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Synergy identification

– EMGs were averaged every 100ms

– The number of synergies was chosen as the minimum number that could explain at least 95% of the variation in the data

Page 9: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Three kicking synergies

Page 10: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.
Page 11: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.
Page 12: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Synergies extracted from jumping swimming and walking

Page 13: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.
Page 14: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Synergy validation

• Are the identified synergies just an arbitrary description of the constraints in the motor output?

• In support of a neural origin of synergies synergy recruitment capture well the pattern

of covariation across different episodes similar synergies are extracted across

behaviors

Page 15: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.
Page 16: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.
Page 17: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.
Page 18: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

______300ms

Page 19: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Summary of results• The muscle patterns recorded in a variety of

natural behaviors can be reconstructed as combination of a small number of muscle synergies

• Synergies are similar across behaviors

• A few synergies are identified only in specific behaviors

• Some synergies have a single dominant muscle and they are part of the same sequence in different behaviors

Page 20: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Focal microstimulation of thelumbar spinal cord hasRevealed a small number of circuits that are organizedto produce muscle synergies.

Page 21: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Motor control primitives in the spinal cord

Mussa-Ivaldi, Giszter and BizziCold Spring Harbor Symposium on Quantitative Biology, vol. 55 (1990)

Page 22: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Regions of the lumbar spinal cord containing the neural circuitry that

specifies the force fields

Page 23: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Tonic Forces

Page 24: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Costimulation of the lumbar interneurons

Page 25: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Motor systems – levels of control

Page 26: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.
Page 27: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.
Page 28: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Examples of Cell Activity Recorded in the Primary Motor Cortex

Page 29: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Two other types of memory cells

Page 30: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Collaborators

A. d’AvellaS. Giszter

F. A. Mussa-IvaldiP. Saltiel

M. TreschVincent C. K. Cheung

Page 31: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

The finding that combination of synergies can explain our data suggest that our synergies may correspond to building blocks of the CPGs, sometimes formulated as a mosaic of “unit burst generators” (Grillner, 1985)

Page 32: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Results• The EMG patterns recorded during

natural motor behaviors can be reconstructed by combinations of a few time-varying muscle synergies

• In some behaviors, there is a systematic relationship between synergy activation coefficients and features of the movement (e.g. kick direction)

Page 33: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Motor systems – levels of control

Page 34: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Copyright ©2005 Society for Neuroscience

Cheung, V. C. K. et al. J. Neurosci. 2005;25:6419-6434

Figure 4. Examples of swimming synergies from analysis stage I

Page 35: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Summary

The main finding is that both intact and deafferented behaviors are primarily generated by the same set of synergies.

Page 36: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Modularity in the spinal cord

• ‘Half-centers’ for the control of rhythmic behaviors (e.g. locomotion) (Brown 1910, Jankowska 1967)

• Central pattern generators (CPGs) by combinations of ‘unit burst generators’ (Grillner 1981)

• Force field modules (Bizzi 1991)

Page 37: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Copyright ©2005 Society for Neuroscience

Cheung, V. C. K. et al. J. Neurosci. 2005;25:6419-6434

Figure 9. Reconstructing the original EMGs with synergies and their coefficients

Page 38: Compositionality - Combinations of Muscle Synergies in the Construction of Motor Behavior Emilio Bizzi Massachusetts Institute of Technology.

Copyright ©2005 Society for Neuroscience

Cheung, V. C. K. et al. J. Neurosci. 2005;25:6419-6434

Stage I analysis of swimming EMGs before and after deafferentation