Mathematical Foundations

Flows on the Plane

Two-dimensional systems — phase portraits, limit cycles, and bifurcations

Dynamics of Living Systems — Theoretical Biology Course

A Brief History

1901
Bendixson — Criterion for closed orbits
Ivar Bendixson established conditions under which closed orbits cannot exist in planar systems, complementing Poincaré’s earlier work and providing the foundation for the Poincaré–Bendixson theorem.
1925
Lotka & Volterra — Predator–prey equations
Alfred Lotka and Vito Volterra independently formulated the first two-dimensional ecological model, predicting oscillations in predator and prey abundances — the canonical example of a planar dynamical system.
1942
Hopf — Bifurcation to periodic orbits
Eberhard Hopf proved that when a pair of complex-conjugate eigenvalues crosses the imaginary axis, a periodic orbit (limit cycle) is born from the equilibrium — the Hopf bifurcation, central to oscillatory dynamics in biology.
1952
Turing — The chemical basis of morphogenesis
Alan Turing showed that a stable homogeneous state in a two-species reaction–diffusion system can become unstable to spatial perturbations, generating patterns. His analysis begins with the 2D Jacobian classification taught in this lecture.
1963
Lorenz — Deterministic chaos requires three dimensions
Edward Lorenz discovered chaotic behaviour in a three-variable model of atmospheric convection. The Poincaré–Bendixson theorem guarantees that such chaos is impossible in two dimensions — making the planar classification (nodes, spirals, limit cycles) complete.
1971
Rosenzweig — Paradox of enrichment
Michael Rosenzweig showed that enriching the prey’s environment destabilises the predator–prey equilibrium via a Hopf bifurcation, creating boom–bust limit cycles — a striking ecological application of the theory developed in this lecture.

Two-Dimensional Systems and Phase Portraits

When two quantities interact — prey and predator, two competing species, cooperators and defectors — the state of the system is a point $(x, y)$ in the phase plane. A two-dimensional autonomous ODE system takes the form:

$$\dot{x} = f(x, y), \qquad \dot{y} = g(x, y)$$

At each point $(x, y)$, the functions $f$ and $g$ assign a velocity vector $(\dot{x}, \dot{y})$. The collection of all such vectors is the vector field. A trajectory is a curve in the phase plane that follows the vector field: starting from an initial condition $(x_0, y_0)$, the system traces a path whose tangent at every point equals the local velocity.

A phase portrait is a picture of representative trajectories, revealing the global dynamics at a glance: where do trajectories converge? spiral? separate? The tools developed in this lecture — linearisation, nullclines, and topological constraints — let us sketch phase portraits without solving the equations.

Linear Systems and the Trace–Determinant Classification

Near a fixed point $(x^*, y^*)$ where $f(x^*, y^*) = g(x^*, y^*) = 0$, the dynamics are governed by the Jacobian matrix:

$$J = \begin{pmatrix} \partial f/\partial x & \partial f/\partial y \\ \partial g/\partial x & \partial g/\partial y \end{pmatrix}\bigg|_{(x^*, y^*)}$$

The linearised system is $\dot{\mathbf{u}} = J\,\mathbf{u}$ where $\mathbf{u} = (x - x^*, y - y^*)$. The eigenvalues of $J$ determine the local behaviour. For a $2 \times 2$ matrix, they depend on only two quantities:

Trace and Determinant

$$\tau = \text{tr}(J) = a + d, \qquad \Delta = \det(J) = ad - bc$$

The eigenvalues are $\lambda_{1,2} = \frac{\tau \pm \sqrt{\tau^2 - 4\Delta}}{2}$.

The entire classification of a 2D fixed point reduces to the $(\tau, \Delta)$ plane:

  • $\Delta < 0$: Saddle. One eigenvalue positive, one negative. Trajectories approach along one direction and diverge along the other.
  • $\Delta > 0$, $\tau < 0$: Stable. Both eigenvalues have negative real part. A node if $\tau^2 > 4\Delta$ (real eigenvalues); a spiral if $\tau^2 < 4\Delta$ (complex eigenvalues).
  • $\Delta > 0$, $\tau > 0$: Unstable. Same distinction between node and spiral, but trajectories diverge.
  • $\Delta > 0$, $\tau = 0$: Centre (in the linear system). Purely imaginary eigenvalues produce closed orbits. In nonlinear systems, this case is delicate: perturbations can turn a centre into a spiral.
  • $\Delta = 0$: Degenerate. At least one zero eigenvalue; a line of fixed points exists.

The parabola $\tau^2 = 4\Delta$ separates nodes from spirals. The $\tau$-axis ($\Delta = 0$) separates saddles from nodes/spirals. The $\Delta$-axis ($\tau = 0$) separates stable from unstable. These three curves partition the $(\tau, \Delta)$ plane into distinct dynamical regimes.

Nullclines and Nonlinear Equilibria

For a nonlinear system, we cannot diagonalise a matrix globally. Instead, we use nullclines: curves where one component of the velocity vanishes.

  • $x$-nullcline: the set where $f(x, y) = 0$. On this curve, $\dot{x} = 0$, so the flow is purely vertical.
  • $y$-nullcline: the set where $g(x, y) = 0$. Here $\dot{y} = 0$ and the flow is purely horizontal.

Fixed points lie at intersections of nullclines — where both components of velocity vanish simultaneously. Between nullclines, the signs of $f$ and $g$ are constant, so the flow direction can be determined by testing a single point in each region. This gives a quick sketch of the global phase portrait.

Once equilibria are located, we classify each by computing the Jacobian $J$ at that point and applying the trace–determinant classification. Linearisation is valid near the fixed point provided neither eigenvalue has zero real part (the Hartman–Grobman theorem).

Conservative Systems and Centres

A system is conservative if there exists a function $H(x, y)$ — a Hamiltonian or first integral — that is constant along trajectories: $\dot{H} = 0$. Trajectories are then confined to level curves of $H$.

The classic example is the Lotka–Volterra predator–prey system (Interacting Species):

$$\dot{N} = rN - aNP, \qquad \dot{P} = eaNP - dP$$

This system admits the conserved quantity $H(N, P) = eaN - d\ln N + aP - r\ln P$. Because $H$ is constant, trajectories form closed loops around the coexistence equilibrium — a centre. The oscillations neither grow nor decay.

Centres are structurally unstable: the slightest perturbation — adding logistic self-limitation to the prey, for instance — breaks the conserved quantity and turns the centre into a stable spiral. This is why the Lotka–Volterra model, despite its elegance, is biologically fragile.

Limit Cycles and the Poincaré–Bendixson Theorem

A limit cycle is an isolated closed trajectory: nearby orbits spiral toward it (stable limit cycle) or away from it (unstable limit cycle). Unlike the closed orbits of a centre, a limit cycle is structurally stable — it persists under small perturbations.

The Poincaré–Bendixson Theorem

If a trajectory of a planar system is confined to a bounded region containing no fixed points, then the trajectory must approach a limit cycle.

This theorem has a profound corollary: chaos is impossible in two-dimensional continuous systems. The long-term behaviour of a planar ODE is restricted to fixed points, limit cycles, or connections between them. Chaos requires at least three dimensions.

To rule out closed orbits, we can use Bendixson's negative criterion: if $\nabla \cdot \mathbf{F} = \partial f/\partial x + \partial g/\partial y$ has one sign throughout a simply connected region, there are no closed orbits in that region. The divergence measures whether the flow expands or contracts area; a closed orbit would require both.

Hopf Bifurcation

The Hopf bifurcation is the birth of a limit cycle from a fixed point. As a parameter $\mu$ varies, a pair of complex conjugate eigenvalues crosses the imaginary axis: their real part changes sign from negative (stable spiral) to positive (unstable spiral), and a limit cycle appears to absorb the trajectories.

The normal form in Cartesian coordinates is:

$$\dot{x} = \mu x - y - x(x^2 + y^2), \qquad \dot{y} = x + \mu y - y(x^2 + y^2)$$

In polar coordinates $(r, \theta)$ this simplifies to $\dot{r} = r(\mu - r^2)$ and $\dot{\theta} = 1$. For $\mu \leq 0$: the origin is a stable spiral ($r \to 0$). For $\mu > 0$: the origin becomes unstable, and a stable limit cycle appears at $r = \sqrt{\mu}$. The amplitude grows as $\sqrt{\mu}$ — a supercritical Hopf bifurcation.

Biological example: the paradox of enrichment (Interacting Species). In the Rosenzweig–MacArthur predator–prey model, increasing the prey's carrying capacity $K$ past a critical value destabilises the coexistence equilibrium via a Hopf bifurcation. The stable spiral becomes an unstable spiral surrounded by a stable limit cycle: the system oscillates with growing amplitude, increasing extinction risk.

In a subcritical Hopf bifurcation, an unstable limit cycle exists before the bifurcation and shrinks onto the fixed point as $\mu$ increases, causing a sudden jump to large-amplitude oscillations or another attractor. This mechanism underlies hysteresis in neural and ecological systems.

Explore: Phase Portraits and the Trace–Determinant Plane

Select a system type. The left panel shows the phase portrait with trajectories from multiple initial conditions. The right panel shows the trace–determinant plane with classification regions; a dot marks the current system.

Phase portrait: trajectories in the $(x, y)$ plane

Trace–determinant plane: classification of equilibria

Exercises

Conceptual Questions

  1. A fixed point has Jacobian with $\tau = -1$ and $\Delta = 3$. Classify the equilibrium and compute the eigenvalues. Will trajectories oscillate as they approach?
  2. Why can a two-dimensional continuous ODE system not exhibit chaos? State the Poincaré–Bendixson theorem and explain which behaviours are possible.
  3. In the Lotka–Volterra predator–prey model, the coexistence equilibrium is a centre. Why is this not structurally stable? What qualitative change occurs when prey self-limitation ($N/K$ term) is added?
  4. Apply Bendixson's negative criterion to the Lotka–Volterra competition model $\dot{N}_1 = r_1 N_1(1 - (N_1 + \alpha_{12}N_2)/K_1)$, $\dot{N}_2 = r_2 N_2(1 - (N_2 + \alpha_{21}N_1)/K_2)$. Compute the divergence $\partial f/\partial N_1 + \partial g/\partial N_2$ in the positive quadrant and determine whether limit cycles are possible.
  5. Explain the Hopf bifurcation in ecological terms. What happens to a predator–prey system when the environment becomes more productive (increasing carrying capacity $K$)?

Computer Problems

  1. Linear Phase Portraits. Implement the 2D linear system $\dot{\mathbf{x}} = A\mathbf{x}$ for the five cases: stable node ($A = \begin{pmatrix} -2 & 0 \\ 0 & -1 \end{pmatrix}$), saddle ($A = \begin{pmatrix} 1 & 0 \\ 0 & -2 \end{pmatrix}$), stable spiral ($A = \begin{pmatrix} -0.5 & 2 \\ -2 & -0.5 \end{pmatrix}$), unstable spiral ($A = \begin{pmatrix} 0.5 & 2 \\ -2 & 0.5 \end{pmatrix}$), and centre ($A = \begin{pmatrix} 0 & 1 \\ -1 & 0 \end{pmatrix}$). For each, integrate trajectories from 8 initial conditions and plot the phase portrait. Compute $\tau$ and $\Delta$ and verify the classification.
  2. Nullcline Analysis. For the Lotka–Volterra competition model with $r_1 = 1$, $r_2 = 0.8$, $K_1 = 100$, $K_2 = 80$, $\alpha_{12} = 0.6$, $\alpha_{21} = 0.8$, plot the nullclines in the $(N_1, N_2)$ plane. Find all equilibria, classify each using the Jacobian, and overlay trajectories from five initial conditions to confirm the global dynamics.
  3. Hopf Bifurcation. Implement the normal form $\dot{x} = \mu x - y - x(x^2 + y^2)$, $\dot{y} = x + \mu y - y(x^2 + y^2)$. Vary $\mu$ from $-0.5$ to $0.5$ in steps of $0.1$. For each $\mu$, plot the phase portrait (trajectories from $r = 0.1$ and $r = 2.0$). Measure the limit-cycle radius for $\mu > 0$ and verify it equals $\sqrt{\mu}$.
  4. Paradox of Enrichment. Using the Rosenzweig–MacArthur model ($r = 1$, $a = 0.5$, $e = 0.3$, $d = 0.3$, $h = 0.5$), vary $K$ from 5 to 25. For each $K$, compute the Jacobian at the coexistence equilibrium, find the eigenvalues, and determine whether the equilibrium is a stable spiral or unstable spiral. Identify the critical $K$ at which the Hopf bifurcation occurs. Plot representative phase portraits before and after the bifurcation.
  5. Trace–Determinant Plane. Write a program that takes any $2 \times 2$ matrix $A = \begin{pmatrix} a & b \\ c & d \end{pmatrix}$ as input, computes $\tau$ and $\Delta$, classifies the equilibrium, plots the phase portrait, and marks the point $(\tau, \Delta)$ on the classification diagram. Test with at least six matrices spanning all regions of the $(\tau, \Delta)$ plane.

References

  1. Bendixson, I. (1901). Sur les courbes définies par des équations différentielles. Acta Mathematica, 24, 1–88.
  2. Hopf, E. (1942). Abzweigung einer periodischen Lösung von einer stationären Lösung eines Differentialsystems. Ber. Math.-Phys. Kl. Sächs. Akad. Wiss. Leipzig, 94, 1–22.
  3. Lorenz, E. N. (1963). Deterministic nonperiodic flow. Journal of the Atmospheric Sciences, 20, 130–141.
  4. Lotka, A. J. (1925). Elements of Physical Biology. Williams & Wilkins.
  5. Rosenzweig, M. L. (1971). Paradox of enrichment: destabilization of exploitation ecosystems in ecological time. Science, 171, 385–387.
  6. Strogatz, S. H. (2015). Nonlinear Dynamics and Chaos (2nd ed.). Westview Press.
  7. Turing, A. M. (1952). The chemical basis of morphogenesis. Philosophical Transactions of the Royal Society of London B, 237, 37–72.
  8. Volterra, V. (1926). Variazioni e fluttuazioni del numero d’individui in specie animali conviventi. Mem. R. Accad. Naz. Lincei, 2, 31–113.