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Inti Suárez *

Zoologisches Institut der Uni6ersität Basel, Rheinsprung 9, CH-4051 Basel, Switzerland

Abstract

Recent developments in dynamical system theory consider chaotic fluctuations of a dynamical system as highly

desirable behaviour because fluctuations allow such a system to be easily controlled. This recent development, chaos

control theory, has been tested in several physical situations. The intrinsic instability of the orbits embedded in a

strange attractor make them easy to change to a more stable and predictable behaviour. The existence of chaos in

natural populations, at least in some species of pest insects, makes the topic of importance in pest management. The

techniques proposed to achieve control over a chaotic dynamical system are presented and discussed in an ecological

context. An important point in the application of this technique to real systems is the amount of noise involved in

the estimation of the parameters of the system. It is also important in the determination of the perturbations required

in the control parameters of the system. The theoretical possibility of obtaining these estimations in field population

biology provides opportunities for using these techniques in pest management. The control of chaos theory shows

how chaotic systems can be controlled by stable and linked systems. The possibility that chaotic behaviour has been

seldom found in nature because of the existence of linked phenomena is discussed. © 1999 Elsevier Science B.V. All

rights reserved.

Keywords: Chaos control; Chaos detection; Pest control; Complex population dynamics

able and advantageous since it has been demon-

Amidst the recent advances in the study of strated that a system sensitive to initial conditions

is highly controllable (Ditto and Pecora, 1993).

chaos, the algorithm developed by Ott et al.

This idea has been explored in an evolutionary

(1990a) has become one of the seminal papers in

context by Doebeli (Doebeli, 1993, 1995a,b) and

the area. It has been said that chaos, considered

in medicine by Weis et al. (1993). The object of

this communication is to explore the conse-

* Tel.: +41-61-2673486; fax: + 41-61-2673457. quences of these developments in population ecol-

E-mail address: suarez@ubaclu.unibas.ch (I. Suárez) ogy, starting from the uses in other contexts.

0304-3800/99/$ - see front matter © 1999 Elsevier Science B.V. All rights reserved.

PII: S 0 3 0 4 - 3 8 0 0 ( 9 9 ) 0 0 0 0 7 - 1

306 I. Suárez / Ecological Modelling 117 (1999) 305–314

1994). They will be discussed later.

The central idea of the method proposed by Ott

et al. (1990a) is to stabilize one of the orbits of the

system’s attractor by making a series of small 3. Chaos control and pest management

perturbations on a control parameter. This is

possible because there is an infinite number of Although the techniques of chaos detection

unstable periodic orbits densely embedded within have been improved (Schaffer, 1984; Sugihara and

the attractor. The proposed algorithm, in general, May, 1990; Ellner and Turchin, 1995), the exis-

is as follows: tence of chaos in natural populations is still con-

1. The system has to be studied through Poincarè troversial, and several reviews about it have been

sections in order to choose a desirable orbit to written (e.g. Hasting et al., 1993). Following

be stabilized. Pimm (1984) the first attempt to detect chaotic

2. With the reconstruction of the attractor in behaviour in nature was important in shifting the

phase space, the direction the system will fol- opinion of ecologists against its existence (Hassell

low in the presence of a determined change in et al., 1976). Nevertheless Hassell et al. (1976) did

the control parameter has to be predicted, find evidence of chaotic behaviour in a pest insect

using the techniques proposed by Lathrop and population, Zeirapthera fagi (Nicholson’s

Kostelich (1989). blowfly). There is some more evidence that it

3. The change in the control parameter that could occur in pest populations (e.g. Phyllaphis

tends to stabilize the desired orbit has to be fagi, Turchin and Taylor, 1992). Then, consider-

made, monitoring the result in phase space, to ing that chaos exists in at least some pest insects,

plan the next change. would it be possible to control these populations

It has been demonstrated than the fast conver- through the control techniques mentioned above?

gence of this algorithm to the desired orbit is a This is the first question that the author intends to

consequence of the instability of the orbits in a answer.

chaotic system (Ott et al., 1990a) and this has Thinking about the applicability of the tech-

been validated in many different experimental sit- niques presented above, one that needs a change

uations (Bayli and Virgin, 1994). in a dynamical variable of the system can be

There are some variations in this method, as compared with a planned harvesting of the popu-

follows: lation (M. Doebeli, personal communication). For

1. Change a dynamic variable from the system, example, to change a dynamical variable in the

not a control parameter from the system. (De- system could be to change the population size as

layed controlling feedback, Pyragas, 1992; dis- convenient (Guemez and Matias, 1993). Because

sipative feedback control, Rulkov et al., 1994). the use of this technique is studied elsewhere

This group of techniques presents less practical (Doebeli and Ruxton, 1997) it will be not dis-

complications than the original one. cussed here. In an ecological context, the main

2. Disturb the control parameter in a periodic advantage of this technique is not needed, because

way, e.g. sinusoidal, (Lima and Pettini, 1990), computer analysis in real time is not a problem,

or with the perturbations in reference to a since the frequency in an ecological system is days

determinate value of the parameter (Hunt, or months, but not nanoseconds. This means that

1991). the human controller can have at least days to

These particular techniques do not need the make the numerical analysis and decide the next

computational resources that the Ott et al. perturbation, whether the system goes on and

(1990a) algorithm does, but their validation is less cycles enough times to allow the next optimal

general. They are more suitable to systems with control. A more relevant point in choosing the

high frequency because they do not need a real appropriate technique is the sensitivity to the

time computer analysis of the system, and they noise present in the determination of the attrac-

I. Suárez / Ecological Modelling 117 (1999) 305–314 307

tor. With regard to this topic Ott et al. (1990b) 1995a). The problem with this kind of manipula-

point out that in the presence of noise, the ampli- tion is in determining exactly what is the magni-

tude of the perturbation that will be made in the tude of the change in the parameter due to the

control parameter must exceed the error length. If controller’s perturbation. If for example, the con-

it is possible to keep the noise bound, it was troller applies some amount of insecticide, he will

shown that the typical outbursts of a chaotic not know immediately the amount of change in

system can be considered as a low probability tail the rate of population growth., but if he main-

case (Ott et al., 1990a), occurring rarely. The tains a monitoring program, he can estimate this

technique proposed by Pyragas (1992) is robust to amount a posteriori, and before making the next

noise. It uses feedback control. If noise is present perturbation.

in the system determination, the feedback in- So far, the feasibility of this kind of control in

creases as the noise increases, achieving control in ecology has been presented. Now, some points

spite of the noise (Pyragas and Tamasevicious, against it will be discussed. Firstly, it must be said

1993). Whatever be the choice between Ott et al.’s that the small perturbations on the system control

(1990a) or Pyragas’s techniques, the possibility of parameters have been permanent in the examples

deciding the amplitudes of the perturbations as a where the statements discussed above were vali-

function of the size of noise needs to be dated. This means that once the parameter is

considered. changed from state p to state p’, the latter will not

To know these amplitudes is, at least theoreti- change to state p’’ or any other until the con-

cally, possible in population field biology. To troller makes the next perturbation. The author

obtain it there are standard techniques to estimate believes that in natural populations it is impossi-

confidence intervals in the determination of popu- ble to affirm the existence of this steadiness.

lation sizes, through capture-recapture methods, Moreover, the opposite cannot be affirmed. The

or another standard technique (Krebs, 1989). Al- resilience of natural populations is still a point of

though a confidence interval, this is not a direct strong debate, with deep implications on manage-

measure of the noise involved in population size ment ecology and conservation (Pimm, 1984).

determination, but it could be used as an indirect Given the disagreements in this debate, the steadi-

measure of it. ness of the control parameter state will not be

Reviewing the proposed techniques, resonant assured until the controlling techniques are ap-

parametric perturbation (Lima and Pettini, 1990) plied to a natural system, and even in that case it

still has to be considered. The control used would will only validate the statements in that particular

have a determined amplitude and frequency, but system and cannot be generalized to other natural

the problem here is that it seems difficult to make systems.

perturbations with well-determined values artifi- Furthermore, the fast convergence of these

cially, in an ecological system. methods has been demonstrated for systems with

Until now there have been presented here some steady changes in the control parameter. Thus,

criteria to choose an adequate technique, and the the convergence speed of the control method re-

possibility of estimating the noise in ecological mains as an open question in ecological systems.

control has been mentioned. The possible exis- Before passing to a numerical example the pro-

tence of a control parameter that could be manip- posed algorithms following from the above argu-

ulated has not been mentioned yet. The logistic ment would be:

model, a system capable of displaying chaotic 1. Obtain historical data on the population to be

behaviour, has only one control parameter: the controlled. With these, build a time series and

population growth rate. It is obvious that this analyze it in phase space, following Schaffer

parameter is susceptible to manipulation, with (1984). It would be useful make additional

biocontrol agents or insecticides. More flexible analyses to find evidence of chaos, following

models, such as the Bellows equation, can also be Ellner and Turchin (1995). Moreover, such an

controlled by perturbing one parameter (Doebeli, analysis permits the derivation of a model of

308 I. Suárez / Ecological Modelling 117 (1999) 305–314

the population dynamics. An estimation of the slope of f at the equilibrium N*, given by f(N*)=

parameters of the model to be used is needed, N*. The slope is given by the first derivative of

to identify stable orbits. f(N) evaluated at N*, N*\0. As this slope

2. Make the perturbation in the control variable, reaches a critical value chaos sets in. The parame-

by, for example, the introduction of an insecti- ter a is not included in the equation for the slope,

cide in the insect’s egg-laying locations. it affects only the equilibrium density, which cor-

3. Follow the amplitude of the change by stan- responds to the carrying capacity of the popula-

dard techniques of estimation of population tion, and will be considered as the control

size. Estimate the size of the noise in this parameter.

determination to plan the size of the next Assuming that parameters l and b have values

perturbation. coding for chaos, it is assumed that it is possible

4. With the new points in the time series, go back to manipulate a in each generation, slightly

to the first step. changing the population ability to cope with the

environment, hence the carrying capacity. In the

case of the control of a pest insect, as discussed

4. A numerical example above, these changes could be reduction of avail-

able egg-laying places, or similar perturbations.

In this section a numerical example of the Monitoring population size, which has a chaotic

control method discussed above is presented. Its dynamic, a generation in which Nt is close to the

efficiency in controlling a biological population equilibrium value, N* will occur. In this genera-

with unstable dynamics will be shown. In the tion, one can approximate the dynamics of the

present case it will be considered that the determi- system linearly:

nation of the population density has a stochastic

(f

component, arising from its estimation in the Nt + 1 − N*= (N ,a ) · (Nt − N*)

wild. The control method is the one proposed by (N t 0

Ott et al. (1990a,b), and it is tried on a simple (f

+ (N*,a0) · (at − a0) (2)

model of population dynamics. The model used (a

was introduced by Maynard-Smith and Slatkin

Introducing the ability to control the change in

(1973), and considered by Bellows (1981) to be the

the parameter a by adjusting it linearly according

most generally applicable, due to its mathematical

to the population density

flexibility. The model and the mathematical con-

siderations for the control of the deterministic al − a0 = c(Nt − N*) (3)

case is briefly described, taken from Doebeli

because c is a constant to be determined, Eq. (3)

(1995a).

could be substituted into Eq. (2), yielding:

Consider a population with discrete generations

whose dynamics are described by: Nt + 1 − N*

Nt + 1 = f(Nt )=Nt

l

(1) =

(f (f n

(N*,a0)+ c (N*,a0) · (Nt − N*) (4)

1+(aNt )b (N (a

Here Nt + 1 and Nt are population densities in If c is chosen to make the modulus of the

successive generations. l, a and b are demo- expression in brackets less than one, the system

graphic parameters: l \ 1 is the intrinsic growth will then approach equilibrium. Evaluating the

) )

rate, a is a measure of how well the individuals partial derivatives, we get the condition

cope with the environment, and b ] 1 describes

l− 1 b(l−1)(b + 1)/b

the type of competition that leads to density 1− b −c B1 (5)

l a 20l

dependence (b: 1 could be interpreted as contest,

b\ \1 to scramble competition). The dynamic The optimal c value occurs when the above

of the population density is determined by the modulus is zero, finally at arriving the equation

I. Suárez / Ecological Modelling 117 (1999) 305–314 309

l −1 a l 2

0

copt = 1−b · (6) with two major perturbations, much smaller than

l b(l − 1)(b + 1)/b

the used in Fig. 1a. If it is intended to control a

To specify when control has to be applied, we natural population with this method, an estima-

must provide a criterion to decide when Nt is close tion of b and l from the real data is needed, and

to N*. As can be seen from Eq. (3), the size of the a time series of 30 or fewer data points is enough

perturbation depends on this decision. A practical to provide estimation of these parameters. In the

strategy to decide the value d for which Nt − simulation shown (Fig. 1b) the parameters of Eq.

N*B dcould be to calculate the size of the maxi- (1) can be estimated from the time series before

mal perturbation that can be done in each the control is activated. Nonlinear regression and

particular system, accordingly calculating d. Con- the Levenberg–Marquardt parameter estimation

sidering that an uncontrolled chaotic population method (provided by SPSS statistical software)

dynamic is ergodic, given any d \ 0 eventually a yield estimation of parameters precise to eight

generation will fulfill the condition (Ott et al., decimal places.

1990b). Fig. 1c shows the system with d= 0.06. The

To include stochastic factors in the simulation, condition to apply the control is only fulfilled

Nt from Eq. (1) is calculated by adding a random after 1100 generations. It is still noteworthy the

percentage of itself: size of the perturbation, which is smaller than in

the preceding cases. Only one relevant perturba-

Ntr

Nt = Nt 9 (7) tion is needed to control the system. In general it

100 is illustrated that the use of smaller values of d

The parameter r is a random number, taken imply a longer time of monitoring the population

from a uniform distribution from zero to a maxi- density to start the control, but once it has been

mal value, fixed at 1, 5, 10 and 20. These extreme made active, the size of the perturbations and its

values are reasonable amounts of noise in the numbers are smaller and the technique works

determination of wild population densities. more efficiently.

(Krebs, 1989). The sign of the added expression Fig. 2 shows the control working with different

was positive with a 0.5 probability each time a levels of noise in the population density. The

population density was calculated, and negative simulations shown are typical of the system, cho-

for the remaining cases. sen after many repetitions.

Fig. 1 shows the effect of choosing different In this case, the technique is robust to signifi-

values of d, namely 10%, 1% and 0.1% of the cant amounts of noise added to the population

maximal population size. In this case no noise in density determination. At increased noise levels,

the population density is simulated. the dynamic is still controlled, but bigger pertur-

Fig. 1a shows the results of the control when bations are needed to keep the system stable.

d = 6 (the maximal population density for the b Even in cases in which the noise is 10% of the

and l values chosen being 60). In the fifth genera- deterministic population density, the method is

tion the control is made active and the equi- still useful for eliminating the fluctuations associ-

librium density is closely attained after three ated with a chaotic regime. These levels of noise

relevant perturbations. The equilibrium orbit is are not enough to force the system out of the

approached asymptotically, so the size of the per- neighborhood of a stable orbit. Only in Fig. 2d,

turbations decreases through time but is not zero. showing a system with 20% noise, is it the case

After the number of perturbations mentioned, the that the stochastic part of a population’s dynamic

changes in them occur at the octave decimal can make the control method no longer useful.

position, and are irrelevant in practice. The reason is that such an amount of noise takes

In Fig. 1b is shown the result of controlling the the system out of the neighborhood of equi-

system when d is set to 0.6. After 30 generations librium. In this case the system returns to chaotic

of uncontrolled chaotic dynamics, the condition is fluctuations. Out of control again, the condition

fulfilled and the population density is controlled to make the perturbation scheme active again

310 I. Suárez / Ecological Modelling 117 (1999) 305–314

Fig. 1. Changing the neighborhood criteria changes the required time for starting the control and the number and size of the

perturbations required. The values of a are shown in the small graphics. The population densities are plotted against generation time

in the main graphics, showing the attainment of the equilibrium density. The parameter values for the figure were as follows:

a0 =0.05, b = 4 and l=5. d= 6 in Fig. 1a, d = 0.6 in Fig. 1b and d= 0.06 in Fig. 1c.

I. Suárez / Ecological Modelling 117 (1999) 305–314 311

Fig. 2. The importance of noise in the control of an unstable dynamic is considered here. The amount of noise increases from Fig.

2a to Fig. 2d. It is noteworthy that in this noisy case, a constant monitoring of the population is needed to keep its dynamics

controlled. As in the previous figure, the a values are shown in the small boxes and the population densities against generation time

in the main graphics. The parameter values for this figure were as follows: a0 =0.05, b =4, l =5 and d=0.6. r =1 in Fig. 2a, r= 5

in Fig. 2b, r= 10 in Fig. 2c and r = 20 in Fig. 2d.

312 I. Suárez / Ecological Modelling 117 (1999) 305–314

takes some time to be fulfilled, but after it hap- ral selection processes). Allen et al. (1993) argue

pens, the system is randomly set out from the that chaotic systems, because of their intrinsic

equilibrium again. noise amplification can amplify the noise of the

This numerical example exemplifies several fea- system, and enhance its natural heterogeneity,

tures of the method mentioned above. It is shown preserving it from extinction.

that knowledge about the levels of noise built into Forgetting the debate about the existence of

the system to be controlled and the maximal size chaos for a moment, let us consider the con-

of the perturbation to be applied are important trolling techniques consisting in the periodic per-

parameters to know before using this control turbation of a main variable of the system (Lima

method properly. It also points to a striking fea- and Pettini, 1990). Now suppose that the periodic

ture of unstable dynamical behavior. To obtain a perturbations occurring in populations, (precisely

stable orbit from a chaotic system, its parameter in those where chaos was expected to be more

does not need to be changed from chaotic values likely, the temperate ones, according to Turchin,

to stable values. Small perturbations at the proper 1993) are the periodic perturbations of a variable

moment are responsible for the change of the of the system. This view can provide a natural

dynamics. For the numerical example shown here chaos control device in an ecological context, but

the changes in the control parameter are displayed there is strong debate about the existence of chaos

in the small windows on the figures. It shows that in the rodent studied (Turchin, 1993, 1995; Falck

the control parameter a is only slightly perturbed. et al., 1995a,b). From this debate, in Turchin’s

In the deterministic case a returns to its original original 1993 work are several reasons to expect

value, but due to the timing of the perturbation chaos in northern populations, but, also in this

the resulting dynamic is stable and not chaotic. northern ecosystem the resources tend to decrease

abruptly during the winter, rising again in the

following seasons, and repeating this pattern in a

5. Chaos control and chaos detection in natural periodic way. Perhaps this seasonal pattern of the

populations: environment can be likened to the parametric

periodic perturbation in the method proposed by

As mentioned at the beginning, the existence of Lima and Pettini. Maybe the non-existence of this

chaos in natural populations is a controversial behaviour in nature is due to periodic, or seasonal

issue. Without dealing with the methodological perturbations, more than the evolutionary argu-

problems for its detection, some arguments ments against chaos, or the non-existence of

against its existence in nature have been pre- proper detection techniques.

sented, by means of natural selection (Berryman This statement, suggested by one of the chaos

and Milstein, 1989). The central argument in this control techniques, agrees with previous works on

work is that if a population behaves chaotically, it metapopulation models. There was controversy

will pass through very small population sizes, on the existence of chaos in connected metapopu-

increasing the probability of extinction, and there- lations (McCallum, 1992; Gonzales-Andujar and

fore cannot be favoured by natural selection. Be- Perry, 1993; Ruxton, 1993, 1996a; Doebeli, 1995a,

sides, it is clearly a group – selectionist argument among others in broader contexts). It hangs on

(and makes invalid the point following Ferriere whether a metapopulation defined as chaotic dis-

and Gatto, 1993, among others), and there are plays chaos when connected (through migration)

models developed that permit us to consider the with another metapopulation whose behaviour

chaotic behaviour of a natural population as an could be, (or not) chaotic. Although there is a

evolutionary stable strategy, the result of natural consensus that it depends on the immigration–

selection acting at the individual level (Ferriere emigration levels, these results are reinforced by

and Fox, 1995, for a review of their views on the the theory developed to explain the synchroniza-

topic; Doebeli, 1993, and Altemberg, 1991 for tion and the control of the chaotic systems (Pec-

alternative views of chaos as an outcome of natu- ora and Carroll, 1990). Furthermore, they are

I. Suárez / Ecological Modelling 117 (1999) 305–314 313

becoming established in a broader theoretical ment and continous discussions about the topic.

frame, the control chaos theory, as pointed out by Two anonymous referees who suggest the addi-

Doebeli (1995b). The fact is that in other disci- tion of a numerical example, improving the read-

plines it has been demonstrated that two poten- ability of the manuscript.

tially chaotic systems, if coupled, can behave in a

periodic fashion, because of their intrinsic chaotic

natures (Pecora and Carroll, 1990). References

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