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Handy Nonlinear Functions for Ecosystem Modeling

Steve Carpenter, Zoology 535

Some common nonlinear functions for ecosystem modeling are presented below. In all
cases, R is the rate, or dependent variable, generally in units of mass/time. S is the substrate, or
independent variable, generally in units of mass. Other terms are parameters. The notation is:

Rmax = maximum rate, mass/time

k = substrate level at which R is half of Rmax, mass

a = dR/dS as S increases a small amount from zero; 1/time

Smax = upper limit of substrate level, mass

q = parameter that controls the slope (dR/dS) near k, for the Hill function

Q = Q10 of a temperature response; the ratio of the rate at 5o + Topt/2 to the rate at Topt/2 - 5o

Topt = optimal temperature

Tmax = upper lethal temperature

All the graphics were generated by the program NonLinDemo.R. Use the program to experiment
with different parameter values, to build intuition about how the parameter values affect the
shapes of the curves.

Asymptotic Relationships

Examples of Asymptotic Relationships

Asymptotic functions: M-M Blue Solid, Exponential Red Dashes
5
4
3
Rate
2
1
0

0 20 40 60 80 100
Substrate
 2

A very common equation for asymptotic relationships is the Michaelis-Menten equation

or rectangular hyperbola:

R = Rmax S / (k + S) (1)

Usually S is a substrate or nutrient concentration, or light intensity, while R is a rate limited by

the substrate, nutrient, or light. In predator-prey interactions, the equation is called a type II
functional response (Holling 1959). S is the prey density, and R is the predation rate. Rmax is the
maximum rate. The parameter k is called the half-saturation coefficient because R is half of Rmax
when S is equal to k.

Another asymptotic relationship with a similar shape is

R = Rmax [1 - exp(-a S / Rmax)] (2)

Here a is the maximum slope of the curve, i.e. the slope when S is near zero. This equation is
sometimes used for the type II functional response in predator-prey interactions, and for the
relationship of photosynthesis rate (R) to light intensity (S).

For examples of additional curves with similar shapes, and a thorough comparison of
light limitation functions for photosynthesis, see Jassby and Platt (1976).

Sigmoid Relationships: The Hill Function

Example of a sigmoid relationship

Hill function
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Rate
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1
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0 20 40 60 80 100
Substrate
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Sigmoid relationships can be modeled by including an additional parameter, q, in the

Michaelis-Menten equation:

R = Rmax Sq / (kq + Sq) (3)

This equation is called the Hill function. Note that when q=1 the Hill function is identical to the
Michaelis-Menten equation. The Hill function is used for the type III functional response in
predator-prey interactions (Holling 1959). Type III predation arises when the predator becomes
more efficient as prey densities increase from very low to moderate values. Learning or
switching behaviors by the predator can lead to this type of predation.

For the original derivation of functional responses for predator-prey systems, see Holling
(1959). A nice explanation of the use of functional responses in ecological models is found in
Hassell (1978). An alternative function for sigmoid relationships, used in studies of animal
behavior and ecological economics, is described by Brock et al. (2006).

Humped Relationships

A simple equation that yields a maximum rate at intermediate values of S is the quadratic
equation

R = Rmax S [1 - (S / Smax)] (4)

This equation is commonly used to describe density-dependent population dynamics, in which R

is the rate of population growth and S is the breeding stock. In this case k is an upper limit to the
population called the carrying capacity. When this quadratic equation is set equal to dS/dt and
integrated, the result is a sigmoid function of time called the logistic equation. Therefore this
rate equation is said to represent logistic population growth.
Quadratic function; note maximum rate
occurs where S is half of Smax.
Logistic
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Rate
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1
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0 20 40 60 80 100
Population Size
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Many populations that exhibit density-dependence do not have maximum growth rates at
S = 0.5*Smax as required by the quadratic equation. A humped curve that can take on
asymmetric shapes is

R = (S / Smax) exp{Rmax [1 - (S / Smax)]} (5)

This equation is called the Ricker equation after the famous fisheries biologist who introduced it
to ecology.

Ricker
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10
Rate
5
0

0 20 40 60 80 100
Spawners

The relationship of organisms' metabolic rates to temperature is also humped. The

maximum rate occurs at an optimal temperature (Topt). Usually the rate is zero at 0oC and at
some upper lethal temperature (Tmax). An additional parameter is necessary to describe the slope
with which the rate rises as temperature increases below Topt. For ecosystem modeling, this
slope Q is defined as the ratio of the rate at 5o + Topt/2 to the rate at Topt/2 - 5o. This definition is
analogous to the Q10 of physiology (Regier and Holmes 1990). With these definitions, the
relationship of temperature to physiological rates is

R = Rmax VX exp[X (1 - V)] where (6)

V = (Tmax - T) / (Tmax - Topt)

X = W2 {1 + [1 + (40/W)]0.5}2 / 400

W = (Tmax - Topt) ln Q

For an example that uses this equation in an ecosystem model, see Bartell et al. (1988). The
equation is sometimes called the Oak Ridge Temperature Function, after the laboratory where it
was first used in ecosystem models.
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Oak Ridge Temperature Function

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Rate
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1
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0 10 20 30 40
Temperature

References

Bartell, S.M., A.L. Brenkert, R.V. O'Neill and R.H. Gardner. 1988. Temporal variation in
regulation of production in a pelagic food web model. pp. 101-118 in: S.R. Carpenter (ed.),
Complex Interactions in Lake Communities. Springer-Verlag, NY.

Brock, W.A., Carpenter, S.R. & Scheffer, M. 2006. Regime shifts, environmental signals,
uncertainty and policy choice. In A Theoretical Framework for Analyzing Social-Ecological
Systems, eds. Norberg, J. and Cumming, G. (Columbia, New York), in press.

Hassell, M.P. 1978. The Dynamics of Arthropod Predator-Prey Systems. Princeton Univ.
Press, Princeton, NJ.

Holling, C.S. 1959. The components of predation as revealed by a study of small predation of
the European pine sawfly. Canadian Entomologist 91: 293-320.

Jassby, A. and T. Platt. 1976. Mathematical formulation of the relationship between

photosynthesis and light for phytoplankton. Limnology and Oceanography 21: 540-547.

Regier, H.A. and J.A. Holmes. 1990. Influence of temperature changes on aquatic ecosystems:
An interpretation of empirical data. Transactions of the American Fisheries Society 119: 374-
389.