← Dynamical Systems in Biology

Jacobian matrices

The Jacobian matrix tells us how the rates of change in a dynamical system respond to small changes in its state variables. It is the multidimensional version of an ordinary derivative.

Core idea. In one dimension, \(f'(x)\) tells us how the rate \(f(x)\) changes when \(x\) changes slightly. In several dimensions, many variables affect many rates, so we need a matrix of derivatives. That matrix is the Jacobian.

Begin with one variable

Suppose

\[\frac{dx}{dt}=f(x).\]

Near a point \(x^*\), a small change \(\Delta x\) produces approximately

\[\Delta f\approx f'(x^*)\Delta x.\]

The derivative therefore measures local sensitivity: how strongly the rate of change responds to a small displacement.

Why one derivative is not enough for two variables

Now suppose

\[\frac{dx}{dt}=f(x,y),\qquad\frac{dy}{dt}=g(x,y).\]

Changing \(x\) can affect both \(dx/dt\) and \(dy/dt\). Changing \(y\) can also affect both rates. We therefore need four local sensitivities.

The Jacobian matrix

\[\boxed{J(x,y)=\begin{pmatrix}\dfrac{\partial f}{\partial x}&\dfrac{\partial f}{\partial y}\\[6pt]\dfrac{\partial g}{\partial x}&\dfrac{\partial g}{\partial y}\end{pmatrix}}.\]

Rows correspond to equations. Columns correspond to variables.

EntryMeaning
\(\partial f/\partial x\)effect of changing \(x\) on the rate of change of \(x\)
\(\partial f/\partial y\)effect of changing \(y\) on the rate of change of \(x\)
\(\partial g/\partial x\)effect of changing \(x\) on the rate of change of \(y\)
\(\partial g/\partial y\)effect of changing \(y\) on the rate of change of \(y\)

How to remember the layout

Take the first differential equation and differentiate its right-hand side with respect to every state variable. That gives the first row. Repeat for the second equation to obtain the second row.

Row = which rate is affected. Column = which state variable is changed.

A simple calculation

Consider

\[\frac{dx}{dt}=x(3-x-y),\] \[\frac{dy}{dt}=y(2-x-y).\]

Write

\[f(x,y)=3x-x^2-xy,\qquad g(x,y)=2y-xy-y^2.\]

The partial derivatives are

\[\frac{\partial f}{\partial x}=3-2x-y,\qquad\frac{\partial f}{\partial y}=-x,\] \[\frac{\partial g}{\partial x}=-y,\qquad\frac{\partial g}{\partial y}=2-x-2y.\]

Therefore

\[\boxed{J(x,y)=\begin{pmatrix}3-2x-y&-x\\-y&2-x-2y\end{pmatrix}}.\]

The Jacobian is usually a function of the state

Notice that this matrix still contains \(x\) and \(y\). Different points in the phase plane generally have different Jacobian matrices.

So there is an important distinction between

\[J(x,y)\]

and the matrix evaluated at a particular equilibrium.

Evaluate at an equilibrium

For the example above, \((3,0)\) is an equilibrium. Substituting \(x=3\), \(y=0\) gives

\[J(3,0)=\begin{pmatrix}-3&-3\\0&-1\end{pmatrix}.\]

This numerical matrix describes the local behaviour near that particular equilibrium.

Why do we evaluate at an equilibrium?

Stability asks what happens after a small displacement from an equilibrium. The Jacobian evaluated there tells us how that small displacement initially changes.

A Jacobian evaluated somewhere else describes local sensitivity there, but it is not directly the standard equilibrium stability calculation.

From a nonlinear system to a local linear system

Let

\[\mathbf{x}^*=\begin{pmatrix}x^*\\y^*\end{pmatrix}\]

be an equilibrium and let a small displacement be

\[\mathbf{u}=\mathbf{x}-\mathbf{x}^*.\]

Near the equilibrium, the nonlinear system can be approximated by

\[\boxed{\frac{d\mathbf{u}}{dt}\approx J(\mathbf{x}^*)\mathbf{u}}.\]

This is called linearisation.

Why linearisation works

A multivariable Taylor expansion gives

\[\mathbf{F}(\mathbf{x}^*+\mathbf{u})\approx\mathbf{F}(\mathbf{x}^*)+J(\mathbf{x}^*)\mathbf{u}.\]

Because \(\mathbf{x}^*\) is an equilibrium,

\[\mathbf{F}(\mathbf{x}^*)=\mathbf{0}.\]

So the leading local behaviour is

\[\mathbf{F}(\mathbf{x}^*+\mathbf{u})\approx J(\mathbf{x}^*)\mathbf{u}.\]
This is why the Jacobian matters: close to an equilibrium, it is the matrix that converts a small displacement into the corresponding approximate rate of change of that displacement.

Geometric intuition

A small displacement \(\mathbf u\) from the equilibrium is transformed by the Jacobian into an approximate local velocity \(J\mathbf u\).

Diagonal entries

The diagonal entries describe local self-effects.

For example, \(\partial f/\partial x\) asks: if \(x\) increases slightly while other variables are held fixed, how does the growth rate of \(x\) respond?

In population models, a negative diagonal term often reflects self-limitation or density dependence, although interpretation always depends on the model.

Off-diagonal entries

The off-diagonal entries describe cross-effects between variables.

For example,

\[\frac{\partial f}{\partial y}\]

measures how changing \(y\) affects the rate of change of \(x\).

These terms can represent competition, predation, infection, mutualism or other biological interactions.

The sign gives local interaction information

If \(\partial f/\partial y\lt0\), increasing \(y\) locally reduces the rate of change of \(x\). If \(\partial f/\partial y\gt0\), increasing \(y\) locally raises it.

But the derivative is local. Its sign and magnitude may change with the state, so it should not automatically be treated as a fixed biological effect everywhere.

Example: predator–prey interactions

Consider

\[\frac{dN}{dt}=rN-aNP,\] \[\frac{dP}{dt}=eaNP-mP.\]

The Jacobian is

\[J(N,P)=\begin{pmatrix}r-aP&-aN\\eaP&eaN-m\end{pmatrix}.\]

The off-diagonal signs have immediate biological meaning:

\[\frac{\partial}{\partial P}\left(\frac{dN}{dt}\right)=-aN\lt0,\]

so more predators reduce prey growth, while

\[\frac{\partial}{\partial N}\left(\frac{dP}{dt}\right)=eaP\gt0,\]

so more prey increase predator growth.

Example: infection coupling

In an epidemic model, an off-diagonal Jacobian entry can measure how a small increase in infectious individuals changes the instantaneous rate of susceptible loss or exposed gain.

This makes the Jacobian useful not only mathematically but also as a compact description of local biological coupling.

From the Jacobian to eigenvalues

Once the Jacobian has been evaluated at an equilibrium, its eigenvalues determine whether small perturbations grow or decay.

If every eigenvalue has negative real part, small disturbances decay and the equilibrium is locally asymptotically stable.

If at least one eigenvalue has positive real part, there is a direction in which disturbances grow, so the equilibrium is unstable.

Why eigenvalues appear

For the linearised system

\[\frac{d\mathbf u}{dt}=J\mathbf u,\]

an eigenvector \(\mathbf v\) satisfies

\[J\mathbf v=\lambda\mathbf v.\]

If a disturbance lies in that direction, its size behaves like

\[e^{\lambda t}.\]

So the eigenvalue tells us whether that particular disturbance mode grows, decays or oscillates.

Trace and determinant in two dimensions

For

\[J=\begin{pmatrix}a&b\\c&d\end{pmatrix},\]

define

\[\operatorname{tr}(J)=a+d,\qquad\det(J)=ad-bc.\]

The eigenvalues satisfy

\[\lambda^2-\operatorname{tr}(J)\lambda+\det(J)=0.\]

Therefore trace and determinant provide a compact route to local stability classification in two-dimensional systems.

Useful two-dimensional rules

ConditionTypical conclusion
\(\det(J)\lt0\)saddle, therefore unstable
\(\det(J)\gt0\), \(\operatorname{tr}(J)\lt0\)both eigenvalues have negative real part: locally asymptotically stable
\(\det(J)\gt0\), \(\operatorname{tr}(J)\gt0\)positive real part: unstable

The discriminant tells us node or spiral

Define

\[D=\operatorname{tr}(J)^2-4\det(J).\]

If \(D\gt0\), the eigenvalues are real. If \(D\lt0\), they are a complex conjugate pair, producing local rotational or oscillatory behaviour.

When linearisation is inconclusive

If an eigenvalue has zero real part, the Jacobian test may not determine the nonlinear stability. Higher-order nonlinear terms may become important.

The Jacobian gives a local first-order approximation. It is extremely useful, but it does not automatically determine global behaviour or every non-hyperbolic equilibrium.

Jacobian versus nullclines

Nullclines show where individual derivatives vanish and divide the phase plane into directional regions. The Jacobian measures local sensitivity and provides the matrix used for local stability analysis.

They answer different but complementary questions.

Jacobian versus the full nonlinear model

The full nonlinear equations describe behaviour throughout the state space. The Jacobian at an equilibrium describes only the leading behaviour close to that point.

Far from the equilibrium, nonlinear effects can dominate and the linear approximation may be poor.

Higher-dimensional systems

For

\[\frac{d\mathbf x}{dt}=\mathbf F(\mathbf x),\qquad \mathbf x=(x_1,\ldots,x_n)^T,\]

the Jacobian is the \(n\times n\) matrix

\[\boxed{J_{ij}=\frac{\partial F_i}{\partial x_j}}.\]

The same principle applies: row \(i\) corresponds to equation \(F_i\), and column \(j\) corresponds to variable \(x_j\).

A practical workflow

Write the differential equations clearly. Differentiate each right-hand side with respect to every state variable. Assemble the Jacobian. Find the equilibria. Substitute each equilibrium into the Jacobian. Calculate eigenvalues, or trace and determinant in two dimensions. Then interpret the result biologically and remember that the conclusion is local unless further analysis establishes global behaviour.

Key idea. The Jacobian is the derivative of a multidimensional dynamical system. It records how every rate responds locally to every state variable. At an equilibrium it gives the linearised system, whose eigenvalues reveal whether small biological perturbations grow, decay or oscillate.