J. theor. Biol. (1994) 166, 275-288 Bifurcation, Chaos and Suppression of Chaos in FitzHugh-Nagumo Nerve Conduction Model Equation S. RAJASEKAR AND M. LAKSHMANAN Department of Physics, Manonmaniam Sundaranar University, Tirunelveli 627 002 and Centre for Non-linear Dynamics, Department of Physics, Bharathidasan University, Tiruchirapalli 620 024, Tamilnadu, India (Received on 19 September 1992, Accepted in revised form on 31 August 1993) We study the effect of constant and periodic membrane currents in neuronal axons described by the FitzHugh-Nagumo equation in its wave form. Linear stability analysis is carried out in the absence of periodic membrane current. Occurrence of chaotic motion, (i) in the absence of both constant and periodic membrane currents, (ji) with constant current only, i) with periodic membrane current only, and (iv) with both constant and periodic currents is investigated for specific parametric choices. We show how chaos sets in through a cascade of period doubling bifurcations. We then demonstrate the ity of control of chaos using various control mechanisms. Specifically, we show the control of possi ‘aos by (adaptive control mechanism, (i) periodic parametric perturbation and (ji) stabilization of unstable periodie orbits. 1. Introduction ‘The propagation of voltage pulse along the mem- branes of the neuronal axons is described by the Hodgkin-Huxley (1952) model. As these equations are highly non-linear, linearizations are not ideal approximations and one would like to have more satisfactory simplified theories. A well-known ap- proximation along these lines is the one suggested by FitzHugh (1961) and used by Nagumo et al. (1962). The coupled partial differential equations (pde) for a voltage pulse V(x,1) along the nerve fibre and a recovery variable R(x, 1) suggested by FitzHugh and ‘Nagumo (see for example, Scott, 1975, 1977) are V.—Vi=FW)ER—1 (ia) R=e(V +a—BR), (ayy where 1 is the membrane current and a, b and c are ‘membrane radius, specific resistivity of the fluid inside the membrane and temperature factor, respectively. Equation (1) describes the spatio-temporal evolution of voltage pulse or excitation variable and recovery (0072-5193/94/030275 + 148.0010 variable or controller variable in nerve membranes. Equation (1) is often called the “cable equation” in electro-physiological applications (Winfree, 1990) where the variable represents local clectric potential across the cell membrane. Controller variable R rep- resents the ionic conductivity of relatively immobile proteins embedded in the cell membrane which act as voltage-sensitive ionic channels, Equation (1) also arises in population genetics (Nagylaki, 1976) where V can be interpreted as the frequency of alleles of one type, 4 say, amongst the total number of alleles in a population of individuals of possible genotypes Ad, a and aa, living in a one-dimensional habitat, When the spatial variation of V, namely, Vy,» is ‘negligible then eqn (1) reduces to a set of two coupled first-order ordinary differential equations (odes), which is known as the Bonhoeffer-van der Pol oscil- fator (Rajasekar & Lakshmanan, 1988a,b, 1992). ‘This two-dimensional oscillator describes local ki tics. On the other hand, secking travelling-wave solutions of the form V= V(x —ut)=V(é) and R= R(x — ut) = R(E), where w is the velocity of the as © 1994 Academic Press Limited216 S. RAJASEKAR AND M. LAKSHMANAN wave, reduces the eqn (1) to the following set of odes (Scott, 1975, 1977): Pa) +R-W 1, 2a) Re -(ch¥ +a -bR), (2b) where dot refers to differentiation with respect to and Jin eqn (1) is assumed to be a function of x — uf. For convenience hereafter we use the notation 1 in place of Numerical evidence (Nagumo et al, 1962; Yamaguti, 1963; Yoshizawa & Kitada 1969; Glendin- ning, 1987) shows that cubic and quadratic non- linearities in F(V) are reasonably adequate forms to account for the shape and the threshold behaviour of nerve pulses. Piece-wise linear form of F(V) has also been considered (McKean, 1970; Rinzel, 1975; Abbott, 1990) in the literature, In recent years, a certain amount of work has been done in the absence ‘of membrane current / for the system (2). In particu- lar, Lakshanan & Rajagopal (1981) have applied a perturbation theory to the system (2) with a= =0 and with cubic non-linearity and obtained Jacobian elliptic function travelling-wave profiles. Casten et al. (1975), Hastings (1976) and Carpenter (1977) have made detailed qualitative analysis in the absence of the external current J. Bursting and relaxation oscit- lations (Tu, 1989) and interaction of pulses (Yamada & Nozaki, 1990) in the absence of external forces have also been studied using a perturbation method. How- ever, from an actual experimental point of view, it is Of great importance to investigate the full non-linear dynamics af the system (2) under the influence of constant and periodic external stimuli as well as the force-free system. In recent times considerable progress has been made in understanding the dynamics of a large class of non-linear oscillators leading to chaotic motion through various bifurcation routes (Holden, 1986; ‘Thompson & Stewart, 1986; Schiister, 1988), thereby ‘opening the possibility of newer understanding of the corresponding physical or biological systems. More interestingly, very recently it has been found that it is possible to control chaos by various algorithmic procedures (Huberman & Lumer, 1990; Lima & Pettini, 1990; Ott et al., 1990; Singer ex al., 199); Rajasekar & Lakshmanan, 1992, 1993), so that the system can be brought back to a desired regular or periodic state with only a minimal effort whenever chaos is found to be undesirable or harmful. ‘The prime aim of the present paper is to investigate the non-linear dynamics of the FN eqn (2) under the influence of external forces as well as the force-frce case and then to study the controlling of chaotic ‘dynamics, In particular, we analyse the effect of the membrane current I= Ag+ Ay cos ot (Qc) jn eqn (2a) with FY) =-V EB. (ea) First, in section 2 we carry out a linear stability analysis for constant /(= 4g). Using combined ana~ lytical and numerical procedures the threshold con- dition for Hopf bifurcation is obtained, We show the existence of a limit cycle motion near the boundary region for specific parameteric choice. In section 3 we numerically investigate the transition from regular to chaotic motion, First, we consider the force-free case, and show that for fixed system parameters, and varying the wave velocity parameter w, the existence of a period doubling route to chaos. Next, fixing the parameter w at a regular regime we discuss the effect of external constant and periodic forces, leading to the onset of chaos. In our study regular and chaotic motions are characterized by using maximal Lya- punov exponent and power spectrum. In section 4 we consider the problem of controlling chaos in the FN equation. We show how adaptive contro! algorithm can be used to set back the system behaviour from a limit eycle to a fixed point and chaotic motion to a period-nT limit cycle motion. The condition for stable control mechanism is derived. Then we study the suppression of chaos by a petiodie parametric pertur- bation, Further, we show the stabilization of an unstable periodic orbit contained in the chaotic at- tractor by slightly adjusting the parameter 4). Finally, section $ contains a summary and con- clusions. 2, Linear Stability Analysis in the Absence of Periodic Membrane Current (J = 49) In this section we wish to investigate the stability of fixed points and the occurrence of Hopf bifur- cation in the absence of periodic membrane current ). Then we discuss the cases of the force-free system (4)=0) and the de driven system (4p #0) separately, For eqn (2), the steady states (fixed points), (Voy Wo, Re), are determined through the equation, V = W(=P)= R =O, that is, through V+ 3¥o(1~b)/b + 3a —bA)Ib=0, Ga) Wi=V,)=0, (Gb) Vy +a —bRy=0. (Gc)BIFURCATION, CHAOS AND SUPPRESSION OF CHAOS am ‘The stability determining eigenvalues are the roots of the characteristic equation P(Q)= B+ a)? +a,4 +4,=0, 4a) where 4, = (u?— be) a= 1— V3 bo, as =c(1 + bV5—b) iu (4b) The stability of a fixed point can be analysed by solving eqn (4). In general, for a fixed point of a dissipative three-dimensional autonomous system to be a stable node all the eigenvalues must be real negative. For a stable focus one of the eigenvalues will bbe negative real and the remaining two must be complex conjugate with negative real part. Thus, a fixed point is asymptotically stable if and only if all the cigenvalues have a negative real part. Hence, the stability can be studied without solving eqn (4) but by determining the sign of the real part of the cigenvalues. According to Routh-Hurwitz’s criterion (Schmidt & Tondl, 1986) a necessary and sufficient condition for a stable stationary solution of (2) to exist is >, a>0, (@4,-a)>0. (40) Equation (4c) is the necessary condition for all. the As to have negative real part. Thus, for a given value of a, b, ¢ and dy a fixed point of (2) is asymptotically stable if and only if eqn (4c) is sat- isfied, namely, >be, Va>max[(b = )/b, (u? = beu?— 6 + b*e?fu"}, (5) where max[4, B] denotes the maximum of A and B. “The steady state may lose its stability when a pair of eigenvalues crosses the imaginary axis with the third eigenvalue being real and negative. Hence, at the threshold values of the parameters, that is, at the Hopf bifureation, P(2) in eqn (4a) must be of the form PA) = + a)(A + BMA — ip) = pal? + PA af? =0, (6a) where a=a>0, P=a>0, af'=a, (6b) From (6b), combining the last two conditions, we obtain a,a,—a=0. Substituting 4, @, as values from eqn (4b) the critical condition for the Hopf bifurcation to occur (a; ~a;= 0) in the system (2) is Vi= be + (6%? chu’, a ‘Now from (3a) the value of 4p at which bifurcation may oceut is given by os = Vox (3 — 3b + BV3.)/(36) + (a/b), (8) where Vo4 are the two roots of V obtained from (7) Now we consider the force free (4 =0) and forcing (4, 0) cases separately, Ase (Di: Ao= 0 Substituting Ao = 0 in (8) the bifurcation values of a for the force-free system are a, = — Voy B36 + V5 2)/3, (a) where Vy = $f) —be + Bie? — cfu” (9b) For 4)=0, we study the Hopf bifurcation in the @—u parameter space. From (9) we note that Vo. is always negative hence either a, or a given by eqn (Ba) becomes negative for a given set of parameters. In a real neural system the negative value of a, the membrane radius, has no biological significance. So wwe consider the case a > 0. In Fig. 1(a) the threshold curve for Hopf bifurcation is plotted in the a-u plane. For systems residing below the locus the stability condition (5) is satisfied and hence the fixed point is stable, Above the threshold curve the fixed point is unstable. However, near the bifurcation value a limit cycle orbit develops due to the Hopf bifurcation. For example, for the choice a = 0-6, 6 = 05, c= 0-1 the system undergoes a Hopf bifurcation at tq = 1-09, Figure 1(b) shows the limit cycle attractor for two different w values chosen close to uy. We will study the bifurcation of this limit eycle in the next section, CASE (11): CONSTANT FORCE (Ap # 0) We now consider the case A, #0. In Fig. 2 we have plotted the bifurcation values of 4) as a function of u for a=05, b=O5 and c=O1l. In the striped region (4,4; ~ a) >0 and the stationary solution is stable. In the remaining region (a, 4; —a;) <0 and the stationary solution is unstable. Limit cycle motion is observed near the bifurcation curve. From Fig. 2 the bifurcation values of 4y for u = 0:9 are doy ©2163 and 4,_ © ~0:163. To know whether the system exhibits limit cycle motion we carried out a detailed ‘numerical study near the bifurcation values. Figures 3 and 4 show the limit cycle attractor for different Ay values chosen close to 4g, and 4y_.. We note that as ‘Ag is decreased (increased) 10 Ao. (do. from higher (lower) values the volume occupied by the limit cycle attractor decreases. On the other hand for the choice w= 0-75a limit cycle is again formed which bifurcates28 S. RAJASEKAR AND M. LAKSHMANAN of Y (a) 3 08 02 ° oS To Ts 2 8 060 ® -070 “tie “in v Fig. |. (a) Bifurcation curve in the a-x plane for a=0'6, b= 05,6 01, 4y=0, 4,=0. Fixed points stable in the sped region. (b) Limit eyle atractor for w= 1406 and | further. The details are again discussed in the next section, 3. Periodic and Chaotic Dynamics In this section we investigate the occurrence of chaotic motion in the FN eqn (2) with (i) 1 =0, (i) T= Ag, (ii) I = A, cos ot and (iv) I = dy + A, 60s ot, (4, #0, A; #0). 30} 20) 10| < 00) 10 20) ° 0308 ~~ O76 «40 Fic. 2. Bifurcation curve in the Ayu plane for a= 05,6 =0'5, =O: and 4, =0, Fixed point is stable in the stiped region. 30 29 & Jer}afa 28 29 a a re v Fig, 3. Limit cycle attractor near the bifurcation value Ap, The da al at a) y= 2104 () 217, (0) Ay 220 td) Aa 3.1. CHAOS IN THE FORCE-FREE SYSTEM (I= 0) We now analyse the response of the system (2) by varying the parameter u, the velocity of the travelling wave solution, for a specific choice of the other parameters. For a= 0:6, b =0°5, c= 0-1 the system hhas only one real fixed point. From Fig. 1 we note that this fixed point is stable for w > 1-09 and unstable for u< 109. At uy, 1-09 the system undergoes Hopf bifurcation and a limit cycle attractor is developed. As the parameter u is decreased from uy, a cascade of period doubling bifurcation of this limit cycle leading to chaotic motion is observed. Figure 5(a) shows the bifurcation phenomenon in the interval u € (1-08, 0-7) where we have plotted the values of (0) intersecting the plane W = V = 0, In this system a simple period- T limit cycle orbit intersects the plane W =0 at two places. Consequently, the Poincaré map consists of ‘two points and will consist of 2n points for orbit of period-nT. A chaotic attractor would exhibit itself & -085 Fic. 4, Limit cycle attractor near the bifurcation value dy_. The Aa values ace (a) dy= 0165, (0) Ay= —017, (6) dy= —03 and 42-023.BIFURCATION, CHAOS AND SUPPRESSION OF CHAOS 29 02 > -07 | Dn EEE EEE Peo eee eee 070 080090 ~«10~«LI0 Fo. 5. Bifurcation diagram (a) and the maximal Lyapunov exponent (b) of the FN model equation for w €(1-08, 07). The other parameters are fixed at =O, b=05, =O, dy=0, A=0. as an infinite number of points in the Poincaré map. From the Fig. S(a) we find that in the tegion we (1-08, 07925) a period-T timit eycle sotution rep- resented by two points in the Poincaré map exists. At 4 =0-7925 bifurcation occurs and a period-27 (four points in the Poincaré map) limit cycle motion devel- ops. As the parameter 1 is decreased further bifur- cations take place and this bifurcation sequence accumulates at u 0-75, One of the most useful quantities to characterize periodic and chaotic attractors is the Lyapunov expo- nents. To estimate the maximal Lyapunov exponent, Ju, WE rewrite eqn (2) in the form of a first-order system X(t) = FOX), ao) Equation (10) is numerically integrated together with the linear equation Y=LXO)YO, ay where L is the linearization of F along the trajectory (0). The maximal Lyapunov exponent is given by dye bm cosa ron) 12) T¥OH, Aa A standard projection of the evolved vectors onto the new orthonormal frame has been used for numerical evaluation of (12). We have calculated the maximal Lyapunov exponent for the FN equation using the Fortran code program given by Wolf et al. (1985). ‘The maximal Lyapunov exponent value against u is plotted in Fig, 5(b). As expected 4, is negative for periodic motion, positive for chaotic regime and is zero at the bifurcation points. Figure 6 shows the chaotic orbit in the (V, 8) plane for u = 0-72. 3.2. CHAOS IN THE DC DRIVEN FN EQUATION (1 = Ag) In this subsection we study the influence of a constant membrane current. Stable fixed point, limit cycle motion and chaotic behaviour are found to occur for a range of values of 4. We illustrate this for specific parametric choice, namely for a =0-5, 6 =0'5, ¢=0'1 and w= 0-75, From eqns (7) and (8) the bifurcation (Hopf) values of y ate 4y, © 2:1094 and do * —0:1094, That is, stationary solution is stable in the region 4, ¢(—0-1094, 2:1094). As shown in section 2, one can expect a stable limit cycle near the bifurcation values. We have numerically analysed the behaviour of the system and we do find the birth of limit cycle oscillations near 4p, and Ay. values, An interesting result is that as the parameter A, is further increased from 4,, or decreased from 4.2 cascade of petiod doubling bifurcations leading to chaotic motion is observed. Figures 7(a) and 8(a) show the bifurcation phenomenon in the interval Ace (—0'11, =0-22) and Aye(2-11,221), Fespect- ively. Here again we have plotted the values of V(0) with 17()=0. The corresponding maximal Lya~ punov exponent is given in Figs 7(b) and 8(b). From the bifurcation diagram 7(a) we find that in the region Age (—0°1094, -0°184) a period-T limit cycle sol- ution (represented by two points in the bifurcation diagram) exists, At A, —0-184 a period-27 timit cycle attractor develops. As the parameter A, is varied further bifurcations take place and this 02 075 00 v in the V-R plane for u =072 6. Chaotic oi280 8. RAJASEKAR AND M. LAKSHMANAN 5 @ 085 Fee wi oo 4-015 ~030 0% O65 moan Ay Fic. 7. Bifurcation diagram (a) and the maximal Lyapunov exponent (b) of the FN oge (2) for Age (011, 0-22) The ather parameters ate fixed at a=0'5, 603, = 0:1 and 4,=0. bifurcation sequence accumulates at Ay —0-199. Chaotic motion appears for 4, €(—0:199, ~0:22), For 4)<—0:22 blow-up of the solution occurs ‘causing one to suppose that either no attractors exist or that the volume of non-divergent points is so attenuated that roundoff errors in computer calculations perturb the orbit out of the basin of attraction. Referring to Fig. 8(a) an infinite sequence of period doubling is observed for 4a (2:1094, 2-199) and chaotic motion is found for 2199 < Ay <2-22, Figure 9(2) and (b) show the chaotic attractor in the (V,R) plane for Ay=—02 and 4y=22, re- spectively. From the above analysis we find that the system exhibits chaos-limit cycle-stable fixed point solution-limit cycle-chaos behaviour when the parameter Ay is increased from —0-22 to 2-22. For Ay>222 blow-up of the solution is found to occur. 3.3. CHAOS IN THE PERIODICALLY DRIVEN FN EQUATION (= 4; COS Of) One of the ways of studying the dynamics of biological oscillators is to subject the oscillators to periodic inputs. In this section we investigate the 080 Ee -10 0-30 — 211 216 eo an a oo 2-045 aa a 7 zit zie 2h 3 os a8 Ae iG, 8, Same as Fig. 7 except for Ay 6 (211,221), Fic. 9. Phase portrait of chaotic attractor for (a) 4y= —0° and. (0) Ay= 22BIFURCATION, CHAOS AND SUPPRESSION OF CHAOS 02 O8 0.05, 00 281 @ = o5 of 06 A © os OF dn 0405; A Fic. 10. Bifurcation diagram (a) and the maximal Lyapunov exponent (6) ofthe FN eqn (2) for 4, (03055). The other parameters ave fixed at a ~05, 6 = 05, 1. 4y~0 and @ =H bifurcation phenomenon in Fig. 10(a). From this figure, itis clear that as 4, is varied from a small value the system undergoes period doubling bifurcations leading to chaotic behaviour. In Fig. 11 we have plotted the chaotic attractor in the (V, R) plane for 4, = 053. The Fourier spectra of the solution of eqn (2) also confirms the period doubling route to chaos. 010. -065 -20 O80 v Fic. 11. Chaotic erbit in the VR plane for A, = 055 The power spectra of a periodic motion with fre- quency w has sharp peaks at w and its harmonics. Chaotic systems have a frequency spectra with a broad-band component. We calculated the power density using a fast Fourier transform algorithm (Press et al., 1986). We have used 2048 data points collected at time interval 2n/(I20). In Fig. 12(a), corresponding to a period-T attractor, the peaks at 0, w/6, w/S alone are clearly visible. The doubling of the number of peaks in Fig. 12(b) and (¢) indicates the period doubling phenomenon. The chaotic case is shown in Fig, 12(d), where the continuous spectrum and the nearby exponential decay for high frequencies supports the chaotic nature of the evolution. The sharp peaks in the continuous spectrum indicates the presence of a large number of unstable periodic orbits im the chaotic attractor. 3.4, EFFECT OF BOTH AC AND DC CURRENTS. (1= Ag+ A, COS ot) In an earlier paper (Rajasekar & Lakshmanan, 19886) we have shown the occurrence of frequency- locking phenomenon in the BVP oscillator by varying282 S. RAJASEKAR AND M, LAKSHMANAN ° @ a) & J : 7 78 2 @ @ | | qe Tio. 12, Power spectrum showing the development of chaos through period doubling bifurcations, The parameter 4 valucs ate 03 (8), 04 (b), 0°55 ©) and 05 (8), the bias 4, and fixing the parameter 4, in a chaotic regime, A winding number diagram consisting of sequence of flat steps, the devils staircase scenario, hhas been obtained. To know whether the FN equation also exhibits mode-locking phenomenon, we have analysed the response of the system for various fixed values of 4, in the interval (0, 0-55) and by increasing the bias 4, step by step. We find that the system does not exhibit frequency locking in this parametric region. However, for a fixed value of 4, as the parameter ‘Ay is varied from a small value, reverse bifurcations leading to period-T limit cyele is found to occur. For example, without 4, as shown earlier in this section, eqn (2) exhibits chaotic motion for 4, =0'55. When 4p is included, the chaotic motion persists for Ay < 00035, For Ay > 0-0035 reverse bifurcation to period-T limit cycle motion is observed. For A, = 05, stable period-4T orbit is found for A, =0 [see Fig. 10(a)]. When 4g is switched on the period-47 orbit persists for 4p <0-0032 and for 4,>0-0032 reverse bifurcation to period-27-speriod-T* oribit_ occurs. The above results clearly show the strong influence of ‘bias 4, on the dynamics on the FN equation. In this section so far we have shown the occurrence of period doubling phenomenon leading to chaotic motion for some sets of parametric values. The FN equation may also exhibit quasiperiodic, intermittent and frequency-locking bebaviours for some other parametric choices. A detailed investigation of the system behaviour in various regions of the parameters space may provide further interesting results. Such a study is in progress. : 4. Controlling of Periodic and Chaotic Motions in the FN Equation Even though deterministic chaos seems to explain many natural processes, in many situations chaos is an undesirable phenomenon which may lead to vio- lent vibrations, irregular operations in mechanical systems, defective functioning of biological systems and s0 on. It is also interesting to note that many cell biologists have been interested in the regular be- haviours, such as symmetric steady-state bend propa~ gation typical of flagella and asymmetric beat cycle with an effective and recovery stroke typical of cilia, but not in the potential irregular behaviours (Segel, 1980). Further, since chaotic motion is sensitive to initital conditions, one naturally wishes to avoid ot reduce chaos so as to improve the performance of the ‘dynamical systems. Moreover, one may notice that a sudden disturbance of a parameter value in a dynam- ical system can lead to a complicated behaviour when the parameter value is shifted to a chaotic regime. It is thus of great practical importance to develop algorithms to contro! chaos and analyse their efficacy. Motivated by the above, in this section we wish to investigate the suppression of, chaos in the FN equation. In particular, we study the controlling of chaos by (i) adaptive control algorithm (ACA) (Huberman & Lumer, 1990), (ii) periodic parametric perturbation and (ii) small feedback contro! mechan- ism (Singer et al., 1991). The basic ideas of these methods are as follows. We consider a general N- dimensional system Xs aN jae = FUG PD, (13a)BIFURCATION, CHAOS AND SUPPRESSION OF CHAOS 283 where X= (%,,%2..+-4%y) are variables and p isa parameter which determines the dynamics of the system. Let X(t) be the chaotic solution of (13a) with specific p(=constant) value. In the ACA, switching the actual motion of the system to a desired state X, can bbe achieved by an additional dynamics on p through (3b) where the function G is proportional to the difference between ¥, and the actual output X and ¢ indicates the stiffness of the control. On the other hand, it is also possible to suppress the chaotic behaviour by means of a small parametric perturbation of suitable fre- quency, that is, by replacing p with p + p'(t), where P(t) is periodic in t, In addition to the above methods, one can suppress the chaotic motion by stabilizing a desired unstable periodic oribit, which will be defined later, embedded in the chaotic orbit. For this purpose, the control parameter p is modified into [p —¢ sgn(¥ ~ 2)]. Here 2 is a desired unstable periodic orbit and sgn is the sign of (X ~ X). In the following we demonstrate the possibility of conver- sion of the actual orbit into a desired orbit using the above three methods for specific choices of par- ameters (as examples). P=GW-X), 4.1. ACA TO THE FN EQUATION WITH Ay = 0, Ay #0 For the FN equation in the absence of periodic ‘component of the membrane current, in order to set back the system motion to a desired state X,=(V,, W,, Ry), We treat do as a control parameter, Under the additional dynamics on 4, the FN equation is written as v=W, (4a) W=-V+VB+R—-uW— 4, (14d) R= ~(clu(V +a—R), (40) —e(V+W+R-V,-W,—R). (14d) For ¢=0, in the absence of control, with a =05, b =05,¢ = 0-1, w= 09, Ay = 2-2 the real fixed point is unstable, However, as noted in section 2 a limit ‘ycle attractor exists due to the Hopf bifurcation. Orbits starting near the unstable fixed point diverge from it and are attracted towards the limit cycle as shown in Fig. 13(a). The initial condition used is (YO), WO), RO) = (08, 0,26). When the control dynamics (14d) are included with 4o(t = 0) = 2:2 and for specific values of ¢ and X, the system evolves under the control dynamics and reaches the steady state X,, This is shown in Fig. 13(b) for ¢=001 and X,= (0817, 0, 2.6355), which is the stable fixed point of the system (I4a-c) for y= 2. 15) a0 @ 10 > 05 ° 100 7200 300 00 t M00 o 19) > 05 ° 300 200 300 400 Fic. 13. Bvolution of variable V of the FN model equation. (8) Limit eyele motion without control (b) Atraction tothe fixed point (@S171,0, 26355) under the contrl for «= 001 Resetting the system motion from chaotic to a limit, cycle using ACA is of great importance, We now consider the conversion of a chaotic motion shown in Fig. (a) for a=05, 6 = 0-5, c= Ol, 4,=0 and A,= 02 into a desired limit cycle orbit and a steady-state (fixed point) attractor. First, we illustrate the conversion of the chaotic motion into a limit cycle. Figure 14(a) shows the chaotic evolution in the V-t plane. The initial conditions used are (¥(0), #(0), R()) =(—08, 0, 08). The parameter 4, evolves according to eqn (144) and adjusts its value until the desired state is reached. Figure 14(b) shows the evolution of the system under the control algor- ithm towards the limit cycle motion, On the other hand, Fig. 14(C) illustrates the conversion of the chaotic flow into a stationary state for ¢ = 0-015 with (VY, Wor R,) = (~0865, 0, ~0-73) which is the stable fixed point of (Ida-c) for 4o= —0-09. In general, the control mechanism is sensitive to € value and the control function G. Thus, itis essential to study the stability of the contro! mechanism, that is, the stability of the desired orbit in the presence of 14(d). The eigenvalue equation in the linear approxi- mation of (14) is given by taht ai taht a= (15a) where a= (= be)ju, a= 1 be — V3 — a, =(1u)(c + bee + beV? — be aec(l +6), ~eu), (5b)284 8. RAJASEKAR AND M. LAKSHMANAN oof =P o > Fae 00a) a w > a5 ° woo 20 300 cgg) e008 A= 008 @ > a 16 ¢ 00 70 woo t Fia. 14. Conversion of chaotic motion (a) to a period-T limit cycle (b) and a stationary state (0 The desired fixed point is thus stable only if a,>0, i=l, 2 3 and 4 (aa-a,)>0 and (a (424, ~ 4,4) — a3) > 0 (Schmidt & Tondl, 1986), Using (15b) these conditions can be simplified into Web, O (be —¢ — be), eal] =u) > W(V3 + Be — +e — 8°, u(t ~u)(be =u) + e[—ute( +5) + ube = 1+ V3) + We (be 1426 —2bY3 — bc) + eu(—b'e +248 + BY?) = be%(1 + be) >(u? whe — WV} Bc? =e) be ~ ¢ —be¥}). (16) For the parametric choice used in Fig. (136), from (16), for stable control we require 0 0 ‘One can also bring the chaotic evolution occurring for d= 0 and 4; > 0 by suitable control dynamics on ,. The FN equation with contro! dynamics on 4 can be written as v=W, (ia) We -V+V3+R-uW—A,coswt, (17b) R= —(clu)(V +a ~bR), (17e) Aye (V+ W+R-V,- WR). =-. cr) In eqn (17) we let 4, evolve as per (17d). Without (17d) for 4 = 0°55 the FN equation exhibits chaotic motion (Fig. 11). Figure 15(a) illustrates the chaotic evolution in the ¥-r plane. We now choose [.), WC), R.(O] as the solution of eqn (I7a-c) with 4, = 03. For this choice of the parameter value a stable period-T limit cycle motion is obsérved. Figure (15b) shows the evolution of the system motion under the control algorithm towards the period-T" limit cycle for ¢=0-1. The attraction towards the period-27 orbit is reported in Fig. 15(0). Further, the dependence of the recovery time Ty, a0 ‘el 00 > 19) ° Ey 300 180~—~—«200 enor i) 00 > ~10 ° 30 100 150 ~~ 200 <=01 © 00 > -10 300180 t Fig, 15, Conversion ofthe chaotic motion (a) toa period-T (b) and petiod-27 (@) limit cyele motionBIFURCATION, CHAOS AND SUPPRESSION OF CRAOS 285 defined as the time taken to reach the desired oribit with the assumed precision 10-4, on cis also studied. For ¢ in the interval (0-01-0-1) we estimated T, for 50 randomly chosen initial conditions and averaged them. As expected 7) decreases when ¢ increases. The applicability of ACA has also been studied using non-linear functions of the form sin 6, @? and exp(¢) in (17d). These non-linear functions have also worked well in bringing back the system to the desired regular motion for a range of control parameter ¢. 4.3, SUPPRESSION OF CHAOS BY PERIODIC PARAMETRIC PERTURBATION Now we study the effect of parametric perturbation added to the FN equation on the chaotic motion. We restricted our analysis to the case 4, > and 4, =0. With periodic parametric perturbation of c, the FN eqn (2) can be written as vew, (18a) We -V4VB+R—uW — A,cosot, (18d) R= ~(Cl +1 cosQnjuy(V ta—bR), 1 <1 (18e) where 1 is the amplitude and Q is the frequency of the perturbation. Starting from a chaotic region (1 =0) we show the suppression of chaos, for ‘example for the case A, =0'55, for some ranges of values of (7,2). For = 0 we tecover the usual FN equation. TABLE | Details of dynamical behaviour of the FN equation in the presence of paramet- ric force for three values of ax a function of n a " Os Obt0s Chaos ‘010 Petiod-16 OO1S-0020 —Period-8 0025-0040 Periods 0085-0050 Chaos 0055-0075 Period 0080-200 ——Period-2 10 ous Chaos ‘0010 Period-16 ‘ots Period:8 0020-0055 ——Peviod-# 0050-0200 Period-2 1S 0000-0025 Chaos 0-030 Period-32 ons Period-16, 0040-0065 ——Period-# 0070-0180 Period 4 0185-0200___Periog-2 00%; 200 A y \ —a-o8 -0075[,~7 O= 10 09 os 34 . Fig. 16. Maximal Lyapunov exponent 4, versus n for = 0'5 ‘and © = 1. The horizontal doted line gives he maximal Lyapunov exponent Z) for 9 =0. We have investigated the behaviour of the system in the (1,9) parameter space. Regular and chaotic motion were detected using the Poincaré map and the Lyapunov exponents. Table 1 summarizes the dynam- ical behaviour of the system (18) as a function of the parameter (0 <1 <0-2) for three fixed values of ©. For 2=0-5 chaotic behaviour persists up to 1 $0005, For 7 >0:005 mode-locked oscillations with periods 16, 8, 4 and 2 are observed. For 2= 1 and 1-5 suppression of chaos by period halving phenomenon is observed as the value of m is increased from zero, Also, we have examined the influence of addition of parametric perturbation to the parameter ‘instead of c. In Fig. 16 we have plotted the maximal Lyapunov exponent (i,) as a function of m for two different © values. The dashed horizontal line rep- resents the 7,, value for = 0. The (7, ©) values with negative 2,, correspond to periodic behaviour. We thus clearly see that for some sets of (n,) values periodic behaviour is recovered. Further, we note that for most of the m values for both = 0°5 and Q= 1, Ant 2) is less than the value Z,,. In other words, even though chaotic behaviour is not suppressed, its strength is reduced, An advantage of this method is that for certain dynamical systems one can study the effect of parametric perturbation on the global bifur- cation using Melnikov analytical technique. Using the periodic parameteric perturbation we can also bri the chaotic evolution occurring for other paramet choices as well, including the cases 4y #0, 4, > 0 to some regular motion. 4.4, STABILIZATION OF UNSTABLE PERIODIC ORBITS It is evident that a chaotic orbit typically has embedded densely within it an infinite number of unstable periodic orbits (UPO) (Auerbach et ai., 1987; Lathrop & Kostelich, 1989). The UPO is defined as follows. Let a chaotic attractor in the Poincaré map of the system (2) be the set of points X1, X35...286 S. RAJASEKAR AND M. LAKSHMANAN, where X =(¥, W, R). Then we call (X,—Xiq4) an UPO of period-k with accuracy 6 if for k >0, Wie KS is satisfied, For example, Fig. 17 shows four different orbits associated with the chaotic attractor of Fig. 1! with 6 =0-05. In addition, chaotic attractors of a system can also contain an unstable steady state (fixed point) fe.g. attractors of the Lorenz system (Sparrow, 1982) and a biochemical network system (Sinha & Ramaswamy, 1987)]. Now we address the following question: given a chaotic system, how to stabilize the UPO or unstable steady state thereby eliminating chaotic behaviour. For this problem, recently, Ott et al, (1990) introduced a method where the par- ameter p in (13a) has been replaced by p + p(t). The determination of p’(1) requires the eigenvalues and eigenvectors of the unstable orbit. However, it is possible to stabilize UPO, without estimating the eigenvalues and eigenvectors, by means of a small feedback control mechanism. In the following we demonstrate this for the FN equation. For an uncontrolled FN system with chaotic motion, first we determine some of the low-period unstable orbits. To stabilize a desired UPO, assuming that the external parameter 4, in eqn (2) is available for small perturbation, we replace A, in (2) by [dy—¢ sen(V — P)]. Here P is the V-component of the unstable period-k oribit, ¢ represents the stiffness of the control signal and sgn corresponds to the sign of (VP), As noted earlier, for 4y=0, ¢=0 the @ o Tan © a -07 -18 os v Fic, 17. Some ofthe unstable period:k orbits embedded in the chaotic attractor for 4, =0°5, o8 @ ~18) ° 80 300 160 200 t 08 © 18, 0 30 Too 160200250 ‘ Fig. 18, Time series plot lustrating the control of an unstable petiod-27 (a) and period-47 (b) orbits, system exhibits chaotic motion for A =0'55. When the controller is switched on at f =f, the controller reacts to the deviation of ¥(¢) from P(t) and modifies the value of 4y. We note that the perturbation takes only either +¢ or —c. We carried out a detailed numerical study for 0 <¢ < 0-2 and for various UPO to observe the effect of the control mechanism. Figures 18(a) and (b) show the conversion of the chaotic motion into stable period-27 and period-T “onbits, respectively, for ¢ = 0-05. 5, Summary and Conclusions In the first part of the present paper using analytical and numerical procedures we have studied the chaotic, dynamics and the effect of constant and periodi membrane currents in the FN equation. The existence of both periodic and chaotic pulses even in the absence of external currents have been pointed out. The system has stable stationary solutions, unstable fixed points, limit cycle motion and chaotic motion depending upon the value of the system parameters. Period doubling route to chaos has been found when the velocity of the travelling wave solution or the amplitude of the external current is varied. In the FN equation the non-periodic solution corresponds to the fact that neurones do not show circadian rhythm while the limit cycte behaviour can explain the pulse trains which code the signal of the single nerve fibres in neural tissue,BIFURCATION, CHAOS AND SUPPRESSION OF CHAOS 287 We wish to mention that in this paper we have studied the dynamics of travelling wave solutions in the FN equation modelled by the system of odes (2). Going back to the original pdes (1), we can interpret the fixed point solutions of (2) as to correspond to the resting state of the voltage pulse. Then limit cycle motion corresponds to the periodic pulse-like sol- utions. Chaotic behaviour in (2) represents chaotic evolution of pulses in a nerve fibre. It is of great importance to know whether or not the solutions of Q) are stable with respect to perturbations in the original partial differential eqn (I). An answer to this question is likely to give some clues to the existence Of patterns in the system. We are currently investi- gating this problem. Further, the bifurcation and chaotic behaviour of travelling wave solutions of (2) reported in this paper may also be amenable to experimental observations. In fact, recently, oceur- rence of period doubling bifurcations leading to chaotic motion (Aihara et al., 1984) and frequency. locking under periodic current have been studied ‘experimentally in squid-axons (Matsumoto et al. 1987). In the second part we considered the problem of controlling of chaos. For specific parametric values we have illustrated the control mechanism by ACA. In the ACA, switching the dynamical nature of the system is by perturbing the system parameter by a suitable rule. The method is sensitive to the stiffness of the control. Before applying the ACA, one should know the system behaviour as a function of the control parameter in order to choose X,. In the periodic parameteric perturbation regular motion is recovered for some ranges of values of the pertur- bation frequency and amplitude. The method dis- cussed in this paper for stabilizing the UPO is ‘comparatively simple and effective. In all the con- trolling procedures, before settling into a desired ‘controlled orbit, the trajectory experiences a chaotic transient and the recovery time is found to be inversely proportional to the stiffaess of the control. Chaotic evolution of voltage pulses in the full FN eqn (1) corresponds possibly to defective functioning of nerve system. In this situation one can control chaos in the real system by a suitable control algor~ ithm. In fact, suppression of chaos in an experimental set up of a Duffing oscillator (Fronzoni et al., 1991) and thermal convection loop (Singer et al., 1991) has been reported recently. Thus, the study of the applica~ bility of various control algorithms in a real nerve system will be of great practical use. Some of the other biological situations where control is believed to play 4 vital rofe include, for example, the maintenance of blood pressure and blood sugar. Clinical experiments on animals showed that the blood pressure is restored to equilibrium values within a few seconds by a sudden perturbation in arterial pressure (Hosomi & Hayashida, 1984). We believe that the control of periodic and chaotic motions explored in section 4 thus has potential utility in such physico-chemical contexts. Further, the conversion to cycles has appli- cability in the regulation of biologically significant ‘oscillatory phenomena ‘The authors thank the referee of this paper for several valuable suggestions. One of us (S.R.) acknowledges M. C. Valsa Kumar for useful discussions On this paper. He also wishes to express his gratitude to the Indira Gandhi Centre for Atomic Research, Kalpakkam for the award of Visiting Scientist Fellowship where part of this work has been done. REFERENCES Aspory, LF. 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