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Exponential Growth
Logistic Growth
Limit cycles, Chaos
Amelioration of chaos
Domain of BIO 201


Ecosystem CHANGING


Population Changed trait values Microevolution

Re-arrangement of genes Speciation
Reduction Division

Enter modeling and the
modeling relationship.
The earliest modeler in ecology

Leonardo Pisano (Fibonacci)

(Born :1170 Died: 1250)
The first law of population dynamics

Nt+1 = Nt + Nt ( b - d + i - e )

Nt = t N0

Why are we calling it a law?

Gives the behavior of a population in the absence of any external

influence (cf HW-law, first law of motion) .

Subject develops by studying departure from the assumptions.

The continuous time analog

= rN

N = N0ert

= er

Does this agree with our observations?

Refining the model: Density dependence

Realized growth rate

= rN (1- N / K)

The continuous logistic equation

Population size

Three possible model outcomes

Population size

Task: Simulate this curve
Empirical verification

Human population size in USA Population dynamics of the rotifer

Brachionus plicatilis
Pearl & Reed 1920
Yoshinaga et al, 2001
Problems with logistic model

The realized growth rate is often not a linearly decreasing function of N.

Realized growth rate


0 200 400 600 800 1000

Hypothetical growth rate function

The Ricker model for dynamics of single population

nt+1= nt exp ( r ( 1 nt / k ) )

Intrinsic growth rate, determines the qualitative nature of

the dynamics

Carrying capacity, the population size at which nt = nt+1

Does not affect the qualitative nature of the dynamics
Behavior of Ricker map
r = 1.3 r = 2.3



Stable point nt 2- point

limit cycle

r = 2.5 r = 3.3


4- point Bifurcation diagram Chaos

limit cycle
The periodicity for a given value of r is equal to the
number of points plotted for that value of r.
Fractals: Self-similarity at multiple scales

Note that this bif diag comes from the logistic model (which looks a bit different from
the Ricker), but the principle of fractals remains the same.

What about the real world?
The Hassell Model (M. P. Hassell, 1975, J. Anim. Ecol. 44: 283-295

Nt+1 = Nt(1+aNt)-
Nt is the population size at time t
is the growth rate parameter
a and are parameters regulating the density dependent feedback term


Hassell et al, 1976, J. Anim. Ecol. 45: 471-486

22/24 exhibited montonic damping.

Conclusion: Most natural populations exhibit relatively simple dynamics

Most simple models of population dynamics predict that populations could, in

principle, exhibit a variety of dynamic behaviors ranging from stable dynamics to
limit cycles to chaos (May, Nature, 1976). This led to a large number of empirical
studies that tried to document chaos in real, biological populations.

Although many populations have very high growth rates, except 1-2 cases, no
real population has been demonstrated to exhibit chaotic dynamics.

Hassell et al, 1976, J. Anim. Ecol. 45: 471- 486 demonstrates that most
populations actually exhibit very simple behavior, which even intuitively does not
make sense.

So, whats going on???????

Making simple models more realistic..
Indivisibility of organisms (Lattice effect)
Henson et al. Science 19 October 2001: 6022

Although real organisms come in integer numbers, population dynamics models do

not incorporate this reality.

So what happens when we impose the condition of integer numbers?

Unintegerized Integerized

Chaos gives way to limit cycles or extinctions

Why does integerisation kill chaos?

Chaos implies that the points in a time series never repeat themselves.
But integerisation, coupled with the upper bound on population size
imposed by the model structure, makes the set of possible values that the
system can take, finite. In other words, sooner or later, a particular value
has to repeat and that is the point where periodicity sets in.

Note that lattice effect can only explain why deterministic chaos can not be
observed in biological systems. It does not exclude the possibility of limit cycles
with long periodicities.
Model complexity
(Turchin and Taylor, Ecology, 73: 289-305 (1992).

Hassell et al. used a model that incorporated only one time lag and a simple
functional form.

Turchin and Taylor fit a large set of models with different number of time lags and
varying complexity

All kinds of behaviors including exponential damping, oscillatory damping, limit

cycles and chaos were observed.

Use of overly simple models for reconstructing
endogenous dynamics from data may be biased in
favor of finding stability.
A visual analogy to understand this

Complex high dimensional objects can have simple low dimensional projections.
So be very careful when you are inferring the former, based on the latter.

Note 1: This is true of life in general.

Note 2: This is just an analogy.
Making models realistic: immigration
McCallum, H. I., J. Theor. Biol., 154, 277284 (1992)

Populations often interact with other populations of the same species, through
exchange of individuals.

Simplest scenario:
What happens when there is a constant immigration of individuals to a population?

nt+1= nt exp ( r ( 1 nt / k ) ) + c

where c is a constant number of individuals that immigrate into the population in

each generation.

Depending on the value of c, chaotic behaviour can be replaced by simple limit cycles

Constant immigration can stabilize the dynamics of populations.


Detailed Reading:
Ch 1 and Ch 2 of Introduction to Population Ecology by Larry Rockwood, Oxford
University Press.
(There is some extra material here which was not covered in the class and
hence will have nothing to do with the exams).

Dey, S. and A. Joshi. Chaos in ecological systems: fact or artifact? Unpublished

(A simple explanation of why integerisation kills chaos)

Extract from Turchin and Taylor, 1992.

(This is a little advanced for this level. You can safely ignore the technical details,
and concentrate on the major conclusions.)

May, R.M. 1976: Simple mathematical models with very complicated dynamics.
Nature 261, 459-467.

The paper that (along with its sibling in 1974) ushered in the era of non-linear

Complex Dynamics in Ecological Time Series

Peter Turchin, Andrew D. Taylor
Ecology, Vol. 73, No. 1 (Feb., 1992), pp. 289-305
(This is a highly cited paper in ecology. Slightly advanced, but people who wish to get
into applied maths or mathematical biology in future are well-advised to grapple with it.)

Extracts from Jurassic Park by Michael Crichton
(Non-technical and very brief expositions on SDIC, fractals and what they all mean in
the context of science. You may or may not agree with all of these, but it is fun to read).