Neural interface dynamics: from spots to spirals Brain and Cortex Principal cells and interneurons Santiago Ramón y Cajal 1900 Eugene Izhikevich 2008 Electroencephalogram (EEG) power spectrum EEG records the activity of ~ 106 pyramidal neurons. Population model Firing rate activity f(E) WEE PE E WEI PI WIE h I WII Ė = Firing rate activity f(I) E + E İ = I I + WEE gEE (A + WII gII (A E) + WEI gEI (A I) + WIE gIE (A+ E) + PE I) + PI WEE PE WEI ⇥(t) = 2 te Q⇥ = E Q= PI WIE I QgjE = f(E) Steady state approximation Qg = f f = f ({g}) ⇥2 t 0 WII 1 d 1+ dt t QgjI = f(I) E = E(gEE , gEI ) g= I = I(gII , gIE ) f Alphoid chaos (10 D) p o w e r frequency PI LLE (S 1 ) inhibition Shilnikov saddle-node route to chaos excitation van Veen and Liley, PRL, 97, 208101 (2006) P Spatially extended models g=w⇥ f Simplest neural field model: Wilson-Cowan (‘72), Amari (‘77) f(u(x, t ⇤ |x ⇤ y|/v)) g = u(x, t) x u(x, t) = w(|x ⇤ y|) dy w(y) y ds (s) f (u(x ⇤ y, t ⇤ s ⇤ |y|/v)) 0 uab = ⇥ab ab 2D layers b ha = uab b wab a p o w e r frequency ab (r, t) = R2 dr wab (r, r )fb hb (r , t ⇤ |r ⇤ r |/vab ) Turing instability analysis E layer and I layer e ik·r e t Continuous spectrum det (D(k, ) ⇤ I) = 0 [D(k, ⇤)]ab = ⇥ab (⇤)Gab (k, ⇤i⇤) = LT G = FLT w(r) (t ⇤ r/v) S Coombes et al., PRE, 76, 05190 (2007) b = f (ss) Re (k) 0.2 = ⇥ + i⇤ 0 2 4 i (k) 6 8 0 6 i 4 6 2 4 0 2 0 a. 20 0 0.2 0 2 15 Turing-Hopf 4 6 10 8 6 c Hopf 5 4 6 2 4 0 0 2 0 b. 0 2 4 6 8 v (axonal speed) 10 12 c Amplitude Equations (one D) Coupled mean-field Ginzburg–Landau equations describing a Turing–Hopf bifurcation with modulation group velocity of O(1). 2 ⇤A1 ⇤ A1 2 2 = A1 (a + b|A1 | + c |A2 | ⇥) + d 2 ⇤⇥ ⇤ + 2 ⇤A2 ⇤ A2 2 2 = A2 (a + b|A2 | + c |A1 | ⇥) + d 2 ⇤⇥ ⇤ Benjamin–Feir (BF) t BF-Eckhaus instability t x k k Coefficients in terms of integral transforms of w and . Applications to co-registered EEG/fMRI Ingo Bojak Bojak, I., Oostendorp, T. F., Reid, A. T., Kotter, R., 2009. Realistic mean field forward predictions for the integration of co-registered EEG/fMRI. BMC Neuroscience 10, L2. A simple 2D neural field model R ut (x, t) = 2 u(x, t) + R2 w(x x )H[u(x , t) 2D Amari model Neural Fields: Theory and Application, (531 pages) Ed. S Coombes, P beim Graben, R Potthast and J J Wright, Springer Verlag, June 2014 h]dx A simulation An interface is easily identified Interface dynamics in 2D n(s) B s r(s) n= n(s ) x u| r(s ) ut (x, t) = t(s ) t(s) x u/| s B u(x, t) + (x, t) (x, t) = w(|x x |)dx B(t) Differentiate u(x, t) = h along B(t) Normal velocity dr u + =0 xu · dt t dr ut n· = dt |z| z ⇥x u(x, t)|x=r ut = dx w(|r h+ ⇥ x |), B zt = z+ dx w(|x x x |) B x=r n = B B K0 - Bessel function of the second kind 2 (1 )K0 (x) = 2⇥ (x) N w(r) = ⇥ ⇥ i r) i=1 dx x w(|x x |) = B dx K0 ( |x B Ai K 0 ( dsn(s)w(|x x (s)|) B x |) = 1 x dsn(s) · |x B r(s) K1 ( |x r(s)| r(s)|) + C 2⇥ 2 Dynamics from data on the boundary only For points on the boundary parametrised by s ⇥ ⇤ N ⌅ ⇥ ut (s) = h + Ai ds n(s ) · Ri (s, s ) + 2 B i=1 zt (s) = ⇤ z(s) Ri (s, s ) = n(s) B s r(s) ds n(s )w(|r(s) r(s )|) B 1 r(s) i |r(s) r(s ) K1 ( r(s )| i |r(s) r(s )|) n(s ) r(s ) t(s ) t(s) i s Biot-Savart style interaction B [effective repulsion between two arc length positions with anti-parallel tangent vectors] Simple numerical scheme Stability of stationary states B0 Zero normal velocity - ut = 0, B(t) ⇥ ⇧ ⇤ N ⌅ ⇥ h= dx w(|r x |) = Ai ds n(s ) · Ri (s, s ) + 2 B0 i=1 B0 i ut = 0 B0 B0 u(t) = u|x B u|x B0 u(B0 ) = h B defines R = R + R(⇥, t) B u(B) = h =0 Spots 2R h = 2⇥ N ⇤ Ai i=1 10 Using Graf’s formula: 1 2 i R ⇥m = m=4 m=3 6 m=3 γ=5 γ=4 4 m=2 m=2 m=2 γ=3 2 0 0 0.05 0.1 K1 ( i R)I0 ( i R) i R(⇥, t) = cos m⇥e m=4 8 R 0.15 h 0.2 1+ ⇥ mt N i=1 Ai Km ( i R)Im ( i R) N i=1 Ai K1 ( i R)I1 ( i R) Stripes (Mex-hat) u(x, y1 ) = h = u(x, y2 ) yi = yi + i cos(kx)e t λ 0.1 0 sinuous -0.1 varicose -0.2 0 0.2 0.4 0.6 0.8 k 1 Linear adaptation 1 ut = u+⇥ at = u ga, a Exploit linearity t t ds (t u(·, t) = ⇥(t) = ⇤ ⇥± = ⇤+ 1+ dse s)⇥(·, s), a(·, t) = +t (1 ⇤+ )e ± (1 + )2 2 Easy to construct stationary spots (1 (t s) ⇤ )e u(·, s) ⇥ t 4 (1 + g) h h(1 + g) Instabilities and travelling pulses Eigenvalues determined by Em ( ) = 0 1 Em (⇥) = (⇥) Wm (1 + g)Wm R = |⇥ (R)| e (⇥) = s (s)ds 0 2 d cos(m )w(R( )) 0 m = 0 mode ( = i⇥ , breath) unstable when g > 1/ emergent frequency ⇥ = m=1 mode ( g 1 R , drift) unstable when g > 1/ Breathing a b c 3 b 2.5 2 0 e a R(t) e d d c 5 10 15 20 25 30 t 35 40 Drifting … at the point where g = 1/ the shape of the spot deviates from circular with an amplitude that depends on quadratic and higher powers of c m R( ) = R + c am cos m m 2 Lu Y, Amari S: Traveling bumps and their collisions in a 2D neural field. Neural Computation 2011, 23:1248–1260 Drifting (weakly nonlinear analysis) For any sigmoid drifting will occur when g increases through 1/ Amplitude analysis (translation operator and drift eigen-modes): 2 X(r, t) = (p) 4S(r) + 2 X j=1 p denotes location of spot ṗ = a 3 aj (t)⇤j (r) + ⇥(r, t)5 a = a1 + ia2 2 ȧ = a(M1 |a| + M2 ) 00 1 000 3 † ⇥M1 = F ⌅1 | ⇤1 ⇥ + F ⌅1 V12 | ⇤†1 ⇥ + ⇧x1 V1 | ⇤†1 ⇥, 6 00 ⇥M2 = F ⌅1 V4 | ⇤†1 ⇥ + (S)⌅1 | ⇤†1 ⇥ + ⇧x1 V4 | ⇤†1 ⇥ spare the details! Scattering Two spots with centers offset by a vector h = 2p 3 ṗ = a + G0 f(p), ȧ = M1 a + M2 a + H0 f(p), f(p) = e z = f(p) -2p 0.1 z t= 0 0.05 3.75 7.5 11.25 / p 15 2p 18.75 22.5 0.3 0 u -0.05 a = velocity -0.1 -0.3 -0.15 0 0.15 a 0.3 −0.1 Spirals Spiral Waves in Disinhibited Mammalian Neocortex (rat slice) Huang et al., J Neurosci. 2004 An interface approach (in progress) Look for rigidly rotating solutions of the form Shape of the spiral arm determined by Stability: … watch this space! In collaboration with Helmut Schmidt Ingo Bojak (Exeter) (Reading) Daniele Avitabile and Aytül Gökçe (Nottingham) The Journal of Mathematical Neuroscience The Journal of Mathematical Neuroscience S Coombes, H Schmidt and I Bojak 2012 Interface dynamics in planar neural field models. EDITORSINCHIEF Stephen Coombes Olivier Faugeras COMING IN 2011 EDITORIAL BOARD Shun-ichi Amari Peter Ashwin Paul Bressloff Romain Brette Bard Ermentrout Wulfram Gerstner Vincent Hakim David Holcman Viktor Jirsa Carlo Laing André Longtin Hinke Osinga Andrey Shilnikov David Terman John Terry Martin Wechselberger AIMS AND SCOPE The Journal of Mathematical Neuroscience (JMN) publishes research articles on the mathematical modeling and analysis of all areas of neuroscience, i.e., the study of the nervous system and its dysfunctions. The focus is on using mathematics as the primary tool for elucidating the fundamental mechanisms responsible for experimentally observed behaviours in neuroscience at all relevant scales, from the molecular world to that of cognition. The aim is to publish work that uses advanced mathematical techniques to illuminate these questions. It publishes full length original papers, rapid communications and review articles. Papers that combine theoretical results supported by convincing numerical experiments are especially encouraged. Papers that introduce and help develop those new pieces of mathematical theory which are likely to be relevant to future studies of the nervous system in general and the human brain in particular are also welcome. Now accepting submissions: www.springer.com/13408