From Lotka-Volterra to mechanism:

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From Lotka-Volterra to mechanism: Simple models have advantages: capturing essential features of dynamical systems with minimal mathematical effort tractable, relatively easy to analyze in full can be parameterized from observation However, they have limited utility: Parameter values are difficult to predict a prior from knowledge of the system

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From Lotka-Volterra to mechanism: . Simple models have advantages: capturing essential features of dynamical systems with minimal mathematical effort tractable, relatively easy to analyze in full can be parameterized from observation However, they have limited utility: - PowerPoint PPT Presentation

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Page 1: From  Lotka-Volterra  to mechanism:

From Lotka-Volterra to mechanism:

Simple models have advantages:

capturing essential features of dynamical systems with minimal mathematical effort

tractable, relatively easy to analyze in full

can be parameterized from observation

However, they have limited utility:

Parameter values are difficult to predict a prior from knowledge of the system

Page 2: From  Lotka-Volterra  to mechanism:

Example:

0 5 10 15 20 250

20

40

60

80

100

120

140

160

Series1Series3

r1 = 0.12K1 = 170a = 0.9 r2 = 0.09K2 = 170b = 0.5

Page 3: From  Lotka-Volterra  to mechanism:

Changes in population size(population dynamics)

Hierarchy of explanation

births migrationsdeaths

Energy balance:food availability

maintenance costcost of reproduction

Risk factors:predator encounters

disease exposurephysical conditions

Behavior:dispersalforaging

group dynamics

Page 4: From  Lotka-Volterra  to mechanism:

Empirical models:

The observations required to estimate parameters are the very same that the model predicts (parameterization = calibrating, fitting).

Population changes

through time

observation

MODEL

parameter estimation

prediction

Page 5: From  Lotka-Volterra  to mechanism:

Mechanistic models:

Some of the observations required to estimate parameters are at least one step removed from the level of prediction.

Population changes

through time

observation

MODELparameter estimation prediction

Page 6: From  Lotka-Volterra  to mechanism:

Tilman’s resource ratio model of plant competition

Observations used to parameterize the model describe resource uptake by plants. Hence, this is a mechanistic model.

loss

Resource level

Biom

ass g

row

th o

r los

s rat

e

growth

loss

Species A

*AR

*AR is the minimal amount

Of resource species A requires to persist in an environment;

If RA is supplied at a certain rate, the species should increase until the resource concentration reaches exactly this value.

Page 7: From  Lotka-Volterra  to mechanism:

Tilman’s resource ratio model of plant competition

loss

Resource level

Biom

ass g

row

th o

r los

s rat

e

growth

loss

Species A

*AR

loss

Resource level

Species B

*BR

When two species are competing for a single limiting resource, the species with the lower equilibrial resource requirement should completely replace the other (B outcompetes A)

Page 8: From  Lotka-Volterra  to mechanism:

Species could be competing for two resources:

loss

Resource 1 level at fixed value of

Resource 2

Biom

ass g

row

th o

r los

s rat

e

growth

loss

Species A

*,1 AR

loss

Resource 2 level at fixed value of

Resource 1

Species A

*,2 AR

Page 9: From  Lotka-Volterra  to mechanism:

Species depend on different resources in different ways:The zero-net-growth-isoclines (ZNG’s)

R1

R2

Resources are perfectly substitutable

R1

R2

Resources are complementary

R1

R2

Resources are perfectly essential

R2*

R1*

Page 10: From  Lotka-Volterra  to mechanism:

Adding resource dynamics

R1

R2

Resource consumption vector

Resourcesupply point; what resources would be without uptake

Resource supplyvector

Page 11: From  Lotka-Volterra  to mechanism:

At equilibrium, both consumers and resources must be unchanging.

Thus, resource supply = resource demand:

R1

R2

Resource consumption vector

Resourcesupply point; what resources would be without uptake

Resource supplyvector

This is where consumer and resource are at equilibrium

Page 12: From  Lotka-Volterra  to mechanism:

Prediction: if different habitats have different resource supply points, resource levels at equilibrium will be

different.

R1

R2

Resource consumption vector

Resourcesupply point; what resources would be without uptake

Resource supplyvector

Page 13: From  Lotka-Volterra  to mechanism:

Species with different resource requirements affect resource levels differently:

R1

R2

R1

Page 14: From  Lotka-Volterra  to mechanism:

What if two species with different resource requirements inhabit the same habitat?

R1

R2

R1

Page 15: From  Lotka-Volterra  to mechanism:

What if two species with different resource requirements inhabit the same habitat?

R1

R2

R1

Page 16: From  Lotka-Volterra  to mechanism:

A two-species equilibrium must be located on both species’ ZNG’s

R1

R2

R1

A and B coexist

A

B

Page 17: From  Lotka-Volterra  to mechanism:

Habitat determines if coexistence is possible.

R1

R2

R1

B wins, because itcan draw R1 to levels intolerable to A.

A

B

Page 18: From  Lotka-Volterra  to mechanism:

R1

R2

R1

A wins, because itcan draw R2 to levels intolerable to B.

A

B

Habitat determines if coexistence is possible.

Page 19: From  Lotka-Volterra  to mechanism:

R1

R2

R1

A wins, because itcan draw R2 to levels intolerable to B.

A

B

Spec

ies B

win

sSpecies A wins

Species A & B coexist

Habitat determines if coexistence is possible.

Both species die

Page 20: From  Lotka-Volterra  to mechanism:

Tilman’s model still predicts the four outcomes of competition that the Lotka-Volterra model does,

and one more: no species lives

AB

A always wins

AB

B always wins

AB

A & B can coexistin some habitats

AB

A & B can coexistin some habitats

B alw

ays w

ins

A always wins

Page 21: From  Lotka-Volterra  to mechanism:

Summary:

What do we gain from Tilman’s more mechanistic model?

Resource requirements for growth can be tested independently of competition.

New predictions: the effect of habitat on species interaction.

Previously overlooked outcomes: both species can fail.

There are predictions we can test and which can fail.

Because the model is process based, we can more easily expand the model to add

more realism.