A quadratic activator can have both basal production and activation inhibited by a second species. A normalized example has reaction rates and , with and positive concentrations. At its spatially homogeneous equilibrium, , so and . Writing , its reaction Jacobian matrix is . Thus and , giving linear asymptotic stability when and instability when . Basal production makes change sign at ; a positive activation feedback needed for a conventional Turing instability requires .
The homogeneous fixed point must satisfy . The singular point is outside the domain of the reaction term, so substitution in the other equation gives the unique regular spatially homogeneous equilibrium
This is the basally forced quadratic activator-inhibitor model. The linearization of its reaction terms gives the Jacobian matrix
Put , so . Its trace and determinant are and , and its eigenvalues are
Since , both eigenvalues have negative real part exactly when . Hence
For the equilibrium is always strictly linearly stable. For , equality is an oscillatory threshold with ; the nonlinear qualification at this threshold is proved below.
Now insert a perturbation proportional to . The diffusion terms subtract from . With the mode trace and determinant are
For a strictly stable homogeneous state, makes for all . A nonzero Fourier mode can therefore grow only if , in which case its two real eigenvalues have opposite signs. The determinant parabola has its minimum at , and that minimum is . The two-species Turing criterion is consequently derived here as
These force , hence . Equivalently, solving for gives
Thus the inhibitor in a reaction-diffusion system must diffuse sufficiently fast. Inside this region, the growing wavenumbers obey
At onset, the minimum touches zero and the two determinant roots coalesce. Therefore
Here is evaluated on the onset surface. These are the Turing threshold of a basally forced quadratic activator-inhibitor model and its continuum critical wavelength. On a finite or domain with periodic boundary conditions an admissible nonzero spatial mode must additionally fall in the stated band. Equality in the diffusion criterion is marginal onset, while is the distinct homogeneous oscillatory threshold.
For completeness, the weak focus at the Hopf threshold of a basal activator-inhibitor model resolves the nonlinear homogeneous stability when . Let , , , and set , , , . Expanding the rational reaction term to cubic order gives
where
Write , and evaluate these homogeneous polynomials on the unit circle. If , , and , division of the radial and angular equations gives
The angular integral of is zero because it changes sign under . Iterating its quadratic term once over a full cycle contributes the square of this zero integral, so the first nonzero return map term is cubic. Using , and the corresponding sixth moments gives and . The Poincaré map is
Because , sufficiently small positive radii decrease on every return. The equilibrium at is therefore nonlinearly asymptotically stable, with weak rather than exponential attraction, despite its neutral linear eigenvalues. The strict inequalities above describe the ordinary diffusion-driven instability from a strictly linearly stable homogeneous state.
First find the positive spatially homogeneous equilibrium. Setting the reactions to zero gives and , and hence the unique positive equilibrium is
The zero pair is not an admissible positive state and is singular in the term. For homogeneous deviations, the reaction Jacobian matrix at this equilibrium is
Thus , , and its eigenvalues are
For , both real parts are negative: a real pair has positive product and negative sum, while a complex pair has real part . The reaction equilibrium is therefore asymptotically stable, with no growing oscillatory mode. This does not exclude damped oscillations. Indeed, for the eigenvalues are complex; for example, gives , a stable focus with oscillatory transients. Thus a literal exclusion of every oscillatory solution is false; the intended exclusion of sustained or growing oscillations is the meaningful statement.
One can strengthen this to exclude nonlinear periodic oscillations in the positive quadrant without relying only on the local linear stability analysis. For the reactions , , use the Dulac function . Then
Were there a periodic orbit wholly in the positive quadrant, the flux of through it would vanish, since this vector is tangent to the orbit; Green's theorem would equate that flux to a strictly negative interior integral. This contradiction proves there is no sustained positive periodic oscillation when , the positive-quadrant Dulac multiplier for quadratic activation argument. It leaves the damped transients entirely consistent with the conclusion.
Now introduce a perturbation in the quadratic activator-inhibitor model. The Fourier mode of squared wavenumber evolves under
Its matrix trace and determinant are
A Turing instability must start with reaction stability, requiring . Then for every , so a mode is unstable exactly when : its two real eigenvalues have opposite signs. This also proves the diffusion-driven onset is stationary rather than oscillatory.
The upward-opening quadratic has its minimum at . To have a negative value at a physically allowed positive squared wavenumber, one needs
Writing , the second inequality is , with roots . The first inequality discards the lower branch, giving
These are the Turing threshold of the quadratic activator-inhibitor model conditions. Equivalently, in the plane the instability region lies strictly above and strictly to the left of , with . The boundary diverges as and approaches as .
Figure 1.
Diffusion-driven instability above the quadratic-model threshold in the reaction-stable parameter strip
. Details.
At equality, the determinant has a double root and one growth rate is zero, while the other remains negative. Substituting into its minimum location gives
Beyond onset, the unstable band is
The stated parameter region assumes continuously available wavenumbers, as on an infinite domain. With specified finite-domain boundary conditions, an allowed nonzero spatial eigenmode must additionally lie in this band; homogeneous stability alone does not ensure that a finite box admits the critical wavelength.
A positive spatially homogeneous equilibrium satisfies and , giving
It exists iff . The reaction Jacobian matrix is
Thus linear stability analysis gives strict homogeneous stability precisely when
The boundaries are not included: at the linearization has purely imaginary exponents, and at the positive equilibrium ceases to exist.
For a mode , put . The perturbation matrix is . In the homogeneously stable region its trace is still negative, so a growing mode exists iff its determinant is negative:
This upward quadratic is negative for some precisely when
Such a Turing instability region is nonempty iff . The unstable band is
For a finite domain only the admissible boundary-condition wavenumbers in this band grow. The region shown below describes a continuous wavenumber spectrum.
Figure 1.
Homogeneous stability and diffusion-driven instability for d=4
. Details.
At onset the minimum of is zero and . On the threshold , substitution gives . Hence
This last expression is the threshold wavenumber; away from onset the fastest-growing mode need not coincide with the minimum of the determinant.
The term makes locally autocatalytic, while increasing suppresses this production. Thus is the activator and is the inhibitor.
At a positive spatially homogeneous equilibrium,
Hence and , giving
The reaction Jacobian matrix there is
Because , the trace-determinant stability criterion shows that the spatially uniform reaction kinetics are stable exactly when
For a perturbation proportional to , put . Its growth rates are the eigenvalues of
The trace is in the reaction-stable regime, while the determinant is
A Turing instability occurs exactly when this upward-opening quadratic is negative for some . By the two-species diffusion-driven instability criterion, this requires
Together these reduce to
Thus the unstable region in the -plane lies below the straight line and to the left of .
Inside this region, the unstable squared-wavenumber band is
At the bifurcation boundary the two roots coincide, so
Substituting yields
At a positive spatially homogeneous equilibrium,
so
The Jacobian matrix of the reaction terms there is
The linear stability analysis of a two-dimensional system requires negative trace and positive determinant. Since , the homogeneous equilibrium is stable exactly when
For a spatial Fourier mode proportional to , put . The linearized matrix is
Its trace is still negative when , while
A Turing instability occurs for some exactly when this quadratic has a negative minimum. This requires
Writing , the second inequality reduces to . Hence, for continuously available wave numbers, the condition is
On a bounded spatial domain, one additionally requires an allowed wave number for which .