An Organizing Center in a Planar Model of Neuronal Excitability
Abstract
This paper studies the excitability properties of a generalized FitzHugh--Nagumo model. The model differs from the classical FitzHugh--Nagumo model in that it accounts for the effect of cooperative gating variables such as activation of calcium currents. Excitability is explored by unfolding a pitchfork bifurcation that is shown to organize five different types of excitability. In addition to the three classical types of neuronal excitability, two novel types are described and distinctly associated to the presence of cooperative variables.
1. , A simple modification of the Hodgkin and Huxley equations explains type 3 excitability in squid giant axons , J. Roy. Soc. Interface , 5 ( 2008 ), pp. 1421 -- 1428 .
2. , In vivo, in vitro, and computational analysis of dendritic calcium currents in thalamic reticular neurons , J. Neurosci. , 16 ( 1996 ), pp. 169 -- 185 .
3. , A novel phase portrait for neuronal excitability , PLoS ONE , 7 ( 2012 ), e41806 .
4. ,
Mathematical Foundations of Neuroscience , Interdisciplinary Applied Mathematics 35 , Springer , New York , 2010 .5. , Geometric singular perturbation theory , J. Differential Equations , 31 ( 1979 ), pp. 53 -- 98 .
6.
R . FitzHugh, Impulses and physiological states in theoretical models of nerve membrane , Biophys. J. , 1 ( 1961 ), pp. 445 -- 466 .7. , Noise-gated encoding of slow inputs by auditory brain stem neurons with a low-threshold K+ current , J. Neurophysiol. , 102 ( 2009 ), pp. 3447 -- 3460 .
8. , The control of firing pattern in nigral dopamine neurons: Burst firing , J. Neurosci. , 4 ( 1984 ), pp. 2877 -- 2890 .
9. , Chattering cells: Superficial pyramidal neurons contributing to the generation of synchronous oscillations in the visual cortex , Science , 274 ( 1996 ), pp. 109 -- 113 .
10. , Slowly varying jump and transition phenomena associated with algebraic bifurcation problems , SIAM J. Appl. Math. , 37 ( 1979 ), pp. 69 -- 106 .
11. , Apamin-sensitive small conductance calcium-activated potassium channels, through their selective coupling to voltage-gated calcium channels, are critical determinants of the precision, pace, and pattern of action potential generation in rat subthalamic nucleus neurons in vitro , J. Neurosci. , 23 ( 2003 ), pp. 7525 -- 7542 .
12. , Membrane bistability in olfactory bulb mitral cells , J. Neurosci. , 21 ( 2001 ), pp. 5311 -- 5320 .
13. , The local electric changes associated with repetitive action in a non-medullated axon , J. Physiol. , 107 ( 1948 ), pp. 165 -- 181 .
14. , A novel T-type current underlies prolonged Ca(2+)-dependent burst firing in GABAergic neurons of rat thalamic reticular nucleus , J. Neurosci. , 12 ( 1992 ), pp. 3804 -- 3817 .
15. , Dynamical Systems in Neuroscience: The Geometry of Excitability and Bursting , MIT Press , Cambridge, MA , 2007 .
16. , Burst firing in dopamine neurons induced by N-methyl-D-aspartate: Role of electrogenic sodium pump , Science , 258 ( 1992 ), pp. 665 -- 667 .
17. ,
Geometric singular perturbation theory , in Dynamical Systems ,Lecture Notes in Math. 1609 , Springer , Berlin , 1995 , pp. 44 -- 120 .18. , Extending geometric singular perturbation theory to nonhyperbolic points---fold and canard points in two dimensions , SIAM J. Math. Anal. , 33 ( 2001 ), pp. 286 -- 314 .
19. , Extending slow manifolds near transcritical and pitchfork singularities , Nonlinearity , 14 ( 2001 ), pp. 1473 -- 1491 .
20. , Relaxation oscillation and canard explosion , J. Differential Equations , 174 ( 2001 ), pp. 312 -- 368 .
21. , Exchange of stabilities in autonomous systems , Stud. Appl. Math. , 54 ( 1975 ), pp. 229 -- 260 .
22. , Exchange lemmas for singular perturbation problems with certain turning points , J. Differential Equations , 167 ( 2000 ), pp. 134 -- 180 .
23. , Sleep and arousal: Thalamocortical mechanisms , Ann. Rev. Neurosci. , 20 ( 1997 ), pp. 185 -- 215 .
24. , A model of the electrophysiological properties of thalamocortical relay neurons , J. Neurophysiol. , 68 ( 1992 ), pp. 1384 -- 1400 .
25. , Different excitability of type 1 and type 2 alpha-motoneurones: The recruitment curve of H- and M-responses in slow and fast muscles of rabbits , J. Neurological Sci. , 28 ( 1976 ), pp. 57 -- 63 .
26. , Voltage oscillations in the barnacle giant muscle fiber , Biophys. J. , 35 ( 1981 ), pp. 193 -- 213 .
27. , Mathematical description of a bursting pacemaker neuron by a modification of the Hodgkin-Huxley equations , Biophys. J. , 16 ( 1976 ), pp. 227 -- 244 .
28. , Asymptotic behavior of solutions of systems of differential equations with a small parameter in the derivatives of highest order , Izv. Ross. Akad. Nauk Ser. Mat. , 21 ( 1957 ), pp. 605 -- 626 .
29. : Insights from simplified membrane models , in Fed. Proc. , Vol. 44 , 1985 , pp. 2944 -- 2946 .
30. ,
Analysis of neural excitability and oscillations , in Methods in Neuronal Modeling , MIT Press , Cambridge, MA , 1989 , pp. 135 -- 169 .31. “type I” excitability in molluscan neurons, in Engineering in Medicine and Biology ,
Proceedings of the 24th Annual Conference and the Annual Fall Meeting of the Biomedical Engineering Society (EMBS/BMES Conference) , Vol. 3 , IEEE,Washington, DC , 2002 , pp. 1966 -- 1967 .32. , Practical Bifurcation and Stability Analysis , 3 rd ed., Interdisciplinary Applied Mathematics 5, Springer-Verlag , New York, 2010 .
33. . Robinson, Threshold firing frequency--current relationships of neurons in rat somatosensory cortex: Type 1 and type 2 dynamics , J. Neurophysiol. , 92 ( 2004 ), pp. 2283 -- 2294 .
34. , A model of a CA3 hippocampal pyramidal neuron incorporating voltage-clamp data on intrinsic conductances , J. Neurophysiol. , 66 ( 1991 ), pp. 635 -- 650 .
35. , Scholarpedia J. , 2 ( 2007 ), p. 1356 .
36. , Up and down states , Scholarpedia J. , 3 ( 2008 ), p. 1410 .
37. , Spontaneous firing patterns of identified spiny neurons in the rat neostriatum , Brain Res. , 220 ( 1981 ), pp. 67 -- 80 .
38. , Current clamp and modeling studies of low-threshold calcium spikes in cells of the cat's lateral geniculate nucleus , J. Neurophysiol. , 81 ( 1999 ), pp. 2360 -- 2373 .