1. Alex Cayco Gajic Journal Club 10/29/13

2. How does motor cortex generate muscle activity? Representational perspective: Muscle activity or abstract trajectory parameters? (e.g. hand velocity) Focus on code in single neurons

3. An epic, twenty-year battle was fought over the cortical representation of movement. Do motor cortex neurons represent the direction of the hand during reaching, or do they represent other features of movement such as joint rotation or muscle output? Graziano 2011 Criticism of the representational approach The role of the motor system is to produce movement, not to describe it. Cisek 2006

4. How does motor cortex generate muscle activity? Representational perspective: Muscle activity or abstract trajectory parameters? (e.g. hand velocity) Focus on code in single neurons Dynamical systems perspective: How can cortex flexibly generate such a large repertoire of movements? Focus on basis sets/ mixed signals, network properties

5. A few equations Representational view: r n (t) = f n (param 1 (t),param 2 (t),…) Dynamical systems perspective: Neural responses muscle movement: m(t) = G[r(t)] But dimensionality of m << dimensionality of r, so G is probably not invertible Dynamical system for population activity: τ r(t) = h(r(t)) + u(t) Dimensionality reduction techniques will be important to find robust, redundant activity patterns

6. Voluntary movements are prepared Random delay period Churchland et al. 2006

7. Voluntary movements are prepared RT decreases with delay period, indicating preparation Churchland et al. 2006

8. Voluntary movements are prepared RT decreases with delay period, indicating preparation Variety of complex single-neuron responses Churchland et al. 2006

9. What is preparatory activity? Representational view Hypothesis: preparatory activity is the subthreshold form of movement activity Churchland et al. 2010a

10. What is preparatory activity? Representational view Hypothesis: preparatory activity is the subthreshold form of movement activity Churchland et al. 2010a

11. What is preparatory activity? Representational view Hypothesis: preparatory activity is the subthreshold form of movement activity Churchland et al. 2010a

12. What is preparatory activity? Representational view Hypothesis: preparatory activity is the subthreshold form of movement activity Reality: preparatory & movement tuning are uncorrelated on average Churchland et al. 2010a

13. What is preparatory activity tuned for? Churchland et al. 2010a Leave out one condition (direction), use linear regression to predict left-out preparatory firing rate from a set of preferred directions. dimensionality PCA analysis: Perimovement: activity of other neurons Kinematic: position, velocity, acceleration EMG: activity for multiple muscles Best performance: from whole population dynamics.

14. What is preparatory activity? Dynamical systems view Hypothesis: preparatory activity brings population dynamical state to an initial value that produces correct motion with minimal reaction time. Churchland et al. 2010b Reduction in variability across different trials as states converge to muscle activation (FF)

15. What is preparatory activity? Dynamical systems view Hypothesis: preparatory activity brings population dynamical state to an initial value that produces correct motion with minimal reaction time. Churchland et al. 2010b

16. What is preparatory activity? Dynamical systems view Hypothesis: preparatory activity brings population dynamical state to an initial value that produces correct motion with minimal reaction time. Churchland et al. 2010b

17. What is preparatory activity? Dynamical systems view Hypothesis: preparatory activity brings population dynamical state to an initial value that produces correct motion with minimal reaction time. Churchland et al. 2010b

18. Convergence of trajectories Reduction in variance comes from convergence of trajectories during motor preparation Churchland et al. 2010b Outlier (monkey hesitated) Covariance ellipses 10-D PCA

19. Convergence of trajectories Reduction in variance comes from convergence of trajectories during motor preparation Churchland et al. 2010b Outlier (monkey hesitated) Prediction: perturbing initial states near go cue should increase RT Covariance ellipses 10-D PCA

20. Preparation & response time Churchland & Shenoy 2007a Use subthreshold microstimulation to perturb prepatory activity No change in wave profile, change in RT only when perturbation occurs at go cue

21. Preparation & response time Use subthreshold microstimulation to perturb prepatory activity No change in wave profile, change in RT only when perturbation occurs at go cue Change in RT due to preparatory state – less dramatic in M1, doesnt exist in saccadic RT Churchland & Shenoy 2007a

22. Preparation & response time Afshar et al 2011 corr(alpha,RT) ?

23. Preparation & response time Afshar et al 2011 Farther along mean neural trajectory smaller RT corr(alpha,RT) < 0

24. PMd neural responses are bizarre Churchland et al. 2010a Tuning differs between preparatory & perimovement periods Response are complex and multiphasic Responses of different neurons are heterogeneous Activity fluctuates longer than movement scale

25. Low-dimensional activity is rotational In low-dimensionality projections, trajectories rotate with phase set by preparatory state (captures ~28% variance) However, the reaches were not overtly rhythmic Brief sinusoidal oscillations form a basis set for more complex patterns Churchland et al. 2012

26. Neural population responses are rotational Churchland et al. 2012

27. Kinematic/EMG data are not Churchland et al. 2012

28. What is jPCA? X = (n)x(ct) matrix PCA to reduce to X red = (k)x(ct) Fit X red = MX red, X red = M skew X red using linear regression V 1, V 2 conjugate eigenvectors of M skew jPC 1 = V 1 +V 2 jPC 2 = j(V 1 -V 2 ) Project X red onto jPC 1, jPC 2 Churchland et al. 2012

29. Conclusions Dynamical systems approach gives insight into movement without making assumptions about single-neuron tuning. Preparation funnels neural trajectories to achieve fast movement without minimal variation. Preparatory state predicts both RT and trial-to-trial variability. Rotational PMd firing rate dynamics form a basis for more complex muscular activity.